The digitized backfile of the magazine of the Arnold Aboretum is browsable here, from its 1911 debut as the Bulletin of Popular Information to its current incarnation as Arnoldia: The Nature of Trees.
Under the “browse by date” button below, click on individual issues to access all articles available in pdf format. Alternatively, click on “browse by author” to access an alphabetical index of contributors with links to their articles.
Electronic access to Arnoldia is embargoed for one year from date of publication, so the most recent issues of the magazine will not be accessible here. Select stories from the most recent issues are posted on the web; see the listing of issues here for links to those articles.
Arnoldia is also made available electronically via JSTOR, accessible through most university and many metropolitan public libraries, and on the web at the Biodiversity Heritage Library.
[{"type":"arnoldia","title":"2022-79-4","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25794","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15e8527.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","arnoldia_cover":true},{"type":"arnoldia","title":"Asian Longhorned Beetle Scouting","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25795","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15e856b.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"12","authors":"Lawlor, Rachel","start_page":"64","end_page":"64","article_content":"Every winter, teams of horticulture staff members scout a random assortment of our susceptible trees for the presence of the invasive Asian long-horned beetle (Anoplophora glabripennis). Maples, elms, willows, horse chestnuts, and poplars are some of their preferred hosts. We record the DBH of the tree, note the start time, and scan the trunk and each branch of a selected tree with binoculars, which is much easier when the trees are lacking foliage. We're looking for any egg-laying sites chewed into the bark by the females, or exit holes from emerging adults after tunneling and feeding on the tree's sapwood and heartwood. A single tree, tall enough and with a spreading crown, could take a group of three over an hour to scout, or a cumulative three work hours to check that specimen. Luckily, ALB has not been detected in the Arboretum. For the health and safety of our collections, we will continue to be vigilant in our yearly scour of the landscape for this destructive pest."},{"type":"arnoldia","title":"Grafting the Past to the Future","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25796","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15e896e.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"11","authors":"Kapplow, Heather","start_page":"61","end_page":"63","article_content":"Though raised on a Pennsylvanian farm, Syracuse-based artist Sam Van Aken was unfamiliar with fruit cultivation when he started working with trees as an art medium. Van Aken's most recent artwork, Open Orchard, is a distributed, site-specific fruit orchard, part of which exists as a long-term art installation on public parkland on Governor's Island in New York City, and the rest of which is scattered around the boroughs of the city, echoing the planting patterns of generations of immigrants that brought plants and seeds with them to New York from their respective homelands. The part of the orchard that is on the island is in an idyllic location, cascading down a gentle slope overlooking New York Harbor. Getting it there was a long journey, beginning in 2014. \\\"I had been shopping around for a place where I could plant one hundred trees in New York City, and people would kind of pat me on the back and say 'good luck!'\\\" Van Aken's lucky break came when he was introduced to curator Shane Brennan, who had just become Director of Public Programs on Governors Island. Brennan responded to the idea immediately, but it was such a drawn-out process that Brennan moved on before Open Orchard was realized. Even finding the right location on the island took several years. The final location was determined by topography. \\\"Fruit trees like to be on raised hills,\\\" Van Aken explains, \\\"so the cold can run off.\\\" But the location ends up also being very resonant with Open Orchard's theme of fruit cultivation as a portrait of the people who have migrated to and lived in New York. \\\"I still pinch myself, because I can't believe that the orchard is there,\\\" he admits. \\\"There's a view when you come from the back part over the hill, and you look down the road, and you see Ellis Island, and the whole project sort of comes together.\\\" The 102 trees on the island will be matched by a roughly equal number planted around New York City, through a collaboration with NYC Parks' community gardening program, Green Thumb (some of these are already in place, others are in a nursery on Governor's Island, in various stages of grafting). Each tree will bear four or five varieties of fruit (apple or stone fruit), all of which are native to or have been historically cultivated in New York. \\\"There are over 100 varieties [of fruit] that originated in New York,\\\" explains Van Aken. Only about two dozen of these are still actively (i.e., commercially) cultivated, but Van Aken was amazed by how much detail he could find about what has grown in New York over the past three hundred years. \\\"I tracked varieties down to intersections, street corners, and farms. And then I took that historical map of New York City, laid it on top of a current one, and realized that I could place these varieties back into the places they originated from. We're currently in the process of doing that. I think today, we've distributed about fifty trees.\\\" Van Aken worked with the U.S. Department of Agriculture (USDA), accessing their germplasm collection to work with varietals that have ceased to exist outside of their archive. \\\"With a fruit tree, you can't [just] store it as a seed, right? So they keep a living sample going. For apples, they store samples in Geneva, New York. For pears, in Corvallis, Oregon. For stone fruit I go to [University of California,] Davis. They'll keep a living example in each of these locations, but then they'll also maintain it as tissue culture.\\\" Trees matched with community gardens get paired with local caretakers, and Van Aken also offers comprehensive grafting workshops and tree-care training materials through Green Thumb as part of the long-term plan for maintaining the offisland part of the orchard. \\\"It's been amazing, distributing trees to people. If you're ever feeling low, just give somebody a tree. I get pictures from the community gardens with five people holding a potted tree and they're just beaming and I'm like, 'I'm doing something right,' you know?\\\" The fruit in Open Orchard spans from pre-contact (pre-colonial) times until about WWII, and includes many fruit varieties that hadn't been accessible for 150-200 years. The newest fruit variety in the orchard is the Stanley plum, developed in Geneva, New York in 1938. \\\"The oldest European variety\\\" says Van Aken, \\\"is the Damson plum. That's believed to date back to Damascus [11 BCE] and was brought into Europe, where it became naturalized and then brought here.\u2026 But, the beach plum, that's been in New York for 10,000 years!\\\" In fact, it may have always been on Governor's Island. \\\"Verrazano [1524] and Hudson [1609] both mentioned it when they sailed into New York Harbor. It's kind of crazy to think that the shoreline of New York was completely lined with these plums that would blossom in white in the spring. It's a far cry from what it [the shoreline] is now.\\\" \\\"When I was looking at all those [tree] placements for community gardens\\\" Van Aken shares, \\\"I realized I had to work with the Lenape Center on this.\\\" Van Aken cold-emailed The Lenape Center, and they enthusiastically invited him into a few of their own projects\u2014for example, an exhibition that the group curated for the Greenpoint branch of the Brooklyn Public Library, which includes a teaching garden composed of \\\"indigenous fruit trees\\\" cultivated by the Lenape in Manhattan, and meant to build \\\"continuity between ecological past and present.\\\" The Lenape Center helped Van Aken arrange for all of the native tree varieties to be planted in Prospect Park (including beach plum, American plum, black cherry, chokecherry, and American persimmon), and also has a future collaboration with Van Aken in the works. Open Orchard grew out of Van Aken's earlier fruit-tree-related work, Tree of 40 Fruit, where the artist pushed the orchardist's practice of grafting to its outer limits. Bearing a title that rings equally biblical and futuristic, Van Aken's Tree of 40 Fruit is a living artwork, with editions spread throughout the world, each producing blossoms and then fruit of 40 different stone fruit varieties. His motivations for constructing Tree of 40 Fruit were among the most traditional artistic drives: he wanted to create something impossible and unexpected\u2014to surprise and delight people when a normal looking tree in their fall\/winter environments burst into multi-colored flower in the spring and then a fruit basket in the summer. But as is often the case in art making, the process became at least as interesting a project as the outcome. In this case, the process brought up big questions about the sustainability of how fruit and knowledge about fruit gets shared. Van Aken's efforts to realize Tree of 40 Fruit raised the specter of varietal extinction. There were once thousands of varieties of stone fruit in the United States, but because of fruit industrialization, it was difficult for him to find and graft scions from enough different varieties for Tree of 40 Fruit to live up to its name. The work thus became an odyssey of archival research and relationship building, and ultimately transformed into a preservation project: purchasers of Tree of 40 Fruit don't just own a unique edition of an artwork. They own an entire cultural history\u2014a living collection of beloved fruits brought from one place to another via immigration\u2014along with some responsibility for keeping rare varieties of peaches, plums, apricots, and cherries present in the future. When searching for fruit varieties for Tree of 40 Fruit, Van Aken found himself welcomed at agricultural research stations and the USDA, but also felt keenly aware that his access to these resources was due to his institutional affiliation (Van Aken is an Associate Professor in the School of Art at Syracuse University). Open Orchard was conceived to make these rare varieties available again for everyone to pick, taste, and modify. He considered a title for the project based on the notion of orchards as commons in Medieval Europe. Then he learned about a philosophy in the Lenape culture \\\"Where essentially, a tree couldn't be owned. You could only come to be gifted the fruit off of the tree.\\\" Open Orchard, with its echo of open-source culture where nothing is proprietary, and anyone can sample and rearrange, clicked into place. \\\"Culture evolves out of agriculture,\\\" Aken says, \\\"and this idea of sharing is an essential component of that, whether that's seeds or knowledge or technology.\\\" The orchard on Governor's Island has been purposely planted with alternating rows\u2014apples every other row, stone fruit in between\u2014so that there can be a (free to the public) harvest within a few years, and so that the apples will be at their peak when the stone fruit trees age out of productivity. But if no one \\\"owns\\\" the trees, who looks out for them? With global warming in mind, irrigation has been installed in Open Orchard, even though orchards don't typically have irrigation. Being on a hill puts the orchard above the flood plains, and the orchard is planted with drought-resistant fescue grass. Van Aken and the island's horticultural team have been working closely on the project since he began grafting the trees for it in their greenhouse 2018, and this close relationship is contractually obligated for at least five years. Van Aken also sees his workshops and community garden tree placements through Green Thumb as part of the orchard's future-proofing process. \\\"Experiencing the fruit is one way of understanding it\u2014aesthetically. But then empowering people through teaching is another way.\\\" Van Aken also describes people becoming excited by reconnecting to their personal histories through caring for their local trees as a strategy for keeping fruit knowledge alive for another generation. Grafting, too, he sees as extremely motivating, almost a mystical experience\u2014a fall graft blossoming in spring still thrills him even though he does about 3,000 grafts a year. \\\"I can talk to people about grafting. I can show them how to graft. I can completely give away all the knowledge. But when that graft takes for that person and they have a thing that they grafted, all of a sudden the mysticism of it comes flooding in.\\\" When meandering among Open Orchard's young trees, imagining them coming into maturity as the view and landscape around them alters due to planned development and unplanned impacts of climate change, it feels as though Van Aken is also grafting in some deeper way\u2014binding forgotten knowledge and biodiversity in place now, to posit a surprising alternative future, branching off from the one we are barreling toward now."},{"type":"arnoldia","title":"City Trees, City Seasons","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25797","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060a326.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"10","authors":"Walker, Kevin","start_page":"58","end_page":"60","article_content":"It is a curious fact that from any of the green public benches found throughout the city of Boston, you can see a tree. And there is no starker contrast to be found in the polarized city of Boston than between the trees of summer and the trees of winter. Throughout Boston's parks and streets, but particularly down along the granite banks of the Charles River, the dreaming trees of summer are as different from the hibernating trees of winter as sunlight is from moon light. The bench where I sit is the bench where I sleep: a green hardwood bench along the Charles River watershed in Boston. My green bench serves double duty as a place to sit and observe the seasons, and, come bedtime, a place for this \\\"rough sleeper\\\" to spend his homeless nights. My bench is in the vicinity of several river trees I've grown to know\u2014and by sleeping between the trees, I know it ain't skid row! I'm in good company with all the gallant elms, sycamores, lindens, mulberry, crabapple, and maple trees found lining the bicycle paths of the Emerald Necklace. And I've noticed over the years camping out that each of these species has a unique reaction to the environment. It's as if a tree could have a personality. It saddens me when a favorite tree loses a large limb due to a harsh winter wind; and I wince in the summer when a large, noble, fully-leaved tree snaps and gets knocked over on top of a footbridge, necessitating its destruction in a giant woodchipper. The occasional stump of a tree looks to me like a boarded up & closed business, reminding me that an important community asset has been lost. Branches stretching skywards towards the light with roots descending into the darkness, the tree is a sentinel of time, a living testament that cannot be recovered if the tree is destroyed by fire or simply knocked over by wind. Silent witnesses to so much Boston history, trees inspire further acts of wisdom: to plant as many trees as possible. Thus, \\\"arborist\\\" is a particularly sagacious vocation. Cheerfully greening and especially beautiful, the trees of summer stretch up towards the sun like so many cathedral spires. Tall and broad like a circus tent, sacred since time immemorial, summer trees are also companionable, providing protection against a sudden rainstorm or providing shade to enjoy a picnic, and glorious, no matter how insect ridden or gnarly they may be. Not all trees are treated equally, however. One particular tree from Asia, called The Tree of Heaven (Ailanthus altissima), is classified as a noxious invasive species. Thought to have been brought to the U.S. in 1784, It is found throughout Boston today. Ailanthus altissima is notorious for producing a chemical that retards the growth of other plants (though other trees do this, too), while its roots can damage sidewalks, sewers, and the foundations of buildings (again, this happens with many other tree species, including natives). No tree is perfect, and I can understand that they sometimes need to be removed. My regret is lessened when I realize that more trees are being planted around Boston on a regular basis. The Charles River estuary is lined with trees, their shiny brass identification tags tinkling in the wind. The early evening darkness is pierced by soft moon glow, silhouetting branches against Boston's crenellated skyline, reminding me of the strange and disturbing paintings of Gustave Dor\u00e9, or an old Civil War daguerreotype, a bleak scene in shades of gray. All trees serve some purpose in the judgement of the arborists who plant them or the neighborhoods that demand them. There are approximately 125,000 trees in the entire city of Boston, and more than 35,000 of these are street trees. Healthy public trees provide much more than shade: they filter air pollution, contribute to climate control, store rainwater and reduce erosion, connect the city to various ecosystems, and provide stable habitat for beneficial insects, birds, and animals. As I limp around the city of Boston, pushing my shopping cart ahead of me in my homeless pilgrimage, the days of tree watching go by slowly, while the years feel quick. I make my way along the footpaths of the Charles River basin, noting the work done on trees. Most of the round brass tags nailed to the trunks have blown off in the winter, while those which remain show various I.D. numbers. Newly planted saplings have plastic tags snapped around their branches, identifying them by species: Northern Red Oak, Swamp White Oak, Little Leaf Linden. I have come to know the trees by the shapes they take, the pattern of their leaves, and the shade or protection they offer. My favorite trees provide deep shade in the searing sun, dense foliage against the down-pouring rain, or wide trunks to offer protection against the sudden winds of a violent squall. Those three types of trees I consider my friends."},{"type":"arnoldia","title":"Olmsted Trees","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25798","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060a36a.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"9","authors":"Greenberg, Stanley","start_page":"50","end_page":"57","article_content":"Much of Brooklyn-based photographer Stanley Greenberg's work explores infrastructure\u2014and as a lifelong New Yorker (and onetime city parksdepartment employee), he knows that parks are urban infrastructure as vital as any aqueduct or reservoir. Greenberg's recent book, Olmsted Trees, documents his hunt for trees surviving from Olmsted's time in parks from New York and Boston to Milwaukee, Wisconsin and Louisville, Kentucky. In a recent conversation, Greenberg reflected on the sometimes awe-inspiring, sometimes unsettling thoughts trees provoke about time and the nature of change. His remarks here are edited and condensed. I 'm in Brooklyn's Prospect Park almost every day. A few years ago I learned that the Prospect Park Alliance had surveyed all the trees in the park, and I realized that I could use it to find the largest and, probably, the oldest trees. With some help from parks staff and an old guidebook, I learned that some of them dated from Olmsted's time. I spent a few days photographing in the park, and then in Olmsted's Chicago parks. Olmsted always said he was designing his parks for how they would look one hundred years from now, and I wondered what the hundred-year-old trees looked like. And that made we wonder if we're thinking enough about one hundred years from our own time. I also realized that Olmsted's two hundredth birthday was coming soon. I decided to try to visit every Olmsted park where he had selected all the plantings. The pandemic slowed my work, but most parks were relatively easy to get to from New York. After many rejections I found a publisher, and my editor wanted a timeline of all the parks in the book, and when the trees were planted. In many parks there were some records, or historical surveys had been done, or arborists showed me their best candidates. I also made educated guesses based on old guidebooks and relative tree sizes. The Arboretum was the only place that had detailed planting records. But as for the timeline, it was almost impossible\u2014because, when is a park finished? Even as far as Olmsted was concerned, he'd work for a few years, and there were probably ten or twenty years of planting to do. So, who's to say when a park is \\\"done\\\"? This is the first project I've done that's been just about \\\"nature.\\\" Much of New York's water supply, which I've photographed extensively, is in somewhat- altered nature. And Olmsted's parks, of course, are mostly constructed (though there were places where he refused to touch what was there). I'm out in nature often, but tend not to photograph when I'm there. This project has shown me that there are things I want to photograph in that world, and has shifted my work in a new direction."},{"type":"arnoldia","title":"Catawba: Back to the Future of the American Wine Industry","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25799","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060a76d.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"8","authors":"Fine, Julia","start_page":"42","end_page":"49","article_content":"In the mid-nineteenth century, American wine drinkers found themselves obsessed with a particular grape, known for its fragrant, somewhat musky profile: 'Catawba', a spontaneously occurring cross between the wine grape Vitis vinifera traditionally cultivated in Europe and the Vitis labrusca grape indigenous to North America's eastern seaboard. By the 1850s, this hybrid had achieved vaunted status in the American wine industry. This was also the time when Ohio was the largest wine-producing state in the country, and approximately 19 out of every 20 vines cultivated there were 'Catawba' grapes.1 Illustrated catalogs of grapes often did not include this \\\"pionee[r] of American grapes\\\" as it was \\\"too generally known to require portrait or illustration by engraving.\\\"2 Even famed poets wrote odes to the grape; Henry Longfellow waxed that this cultivar \\\"has a taste more divine, \/ more dulcet, delicious and dream\\\" than any other, so much so that it \\\"has need of no sign, \/ No tavern-bush to proclaim it\\\" as it was so eminently popular.3 'Catawba'\u2014which American winemakers often argued distinguished their wine from European classics\u2014was thus the pi\u00e8ce de r\u00e9sistance of the early American wine industry. It may seem surprising, then, that this grape fell almost entirely out of favor in the post-Prohibition era, surpassed by classic V. vinifera grapes. Today, few Americans have heard of 'Catawba'. Those who have may associate it with saccharine pink fortified and unfortified wines and grape juices. Across the United States, many sommeliers who are familiar with 'Catawba' decry it as \\\"foxy,\\\" \\\"musky,\\\" or \\\"unidimensional,\\\" and eschew wines made with it.4 Why was a widely popular grape relegated to the realm of the outmoded, even unknown? Through seed catalogues, regional publications, and viticulturists' correspondence, we can trace the rise and fall of 'Catawba' and the Ohio wine industry which is often ignored in contemporary scholarship in favor of the later emergence of the California wine industry. But, as environmental historian Richard White reminds us, \\\"in paths forged and blocked, abandoned and resumed, history shows that things need not be the way they are.\\\"5 Once shunned by the American palate, the 'Catawba' has the potential to contribute to the reinvigoration of the American wine industry in the face of the climate crisis. Towards a Botanical History of 'Catawba' First, though, what exactly is 'Catawba'? The history of the grape, much like its parentage, is murky. There are many stories regarding how settler winemakers came across the grape in the nascent United States. Some historians (perhaps apocryphally) suggest that one \\\"General Davy\\\" living on the Catawba River brought the cultivar to Washington DC and distributed it among friends in that area, associating the cultivar with the river (which was itself named after the Catawba or Iswa Indigenous people).6 While the veracity of this story is unclear, we know that by 1823, 'Catawba' became more widespread, helmed by the \\\"Father of American Viticulture\\\" Major John Adlum who promoted indigenous grape varietals and attempted to create viticultural experiment stations. In an 1829 letter reprinted in the New England Farmer, and Horticultural Register, Adlum reported how he discovered a 'Catawba' vine in \\\"Mrs. Schell's garden,\\\" in Montgomery County, Maryland. While Mrs. Schell apparently had no information as to how the family acquired this vine, she informed Adlum that her husband had referred to it as the Catawba grape.7 Adlum, for his part, sent cuttings of the plant around the country, including to Ohio viticulturist Nicholas Longworth, who, as we will see, played a critical role in popularizing the cultivar. From its beginnings, viticulturists have argued over the exact botanical definition of 'Catawba', with some claiming it was entirely a variety of Vitis labrusca, some contending it was V. labrusca mixed with another indigenous American grape species, and others still arguing that it was a mix between V. labrusca and V. vinifera.8 Today, it seems as if the last hypothesis is correct, as most present-day viticulturists believe 'Catawba' is a spontaneously occurring hybrid between labrusca and vinifera that emerged after European settlers brought cuttings of V. vinifera to North America.9 In particular, Catawba is probably a hybrid between V. abrusca grapes and \\\"S\u00e9millon,\\\" a white V. vinifera grape commonly grown in the southern winemaking regions of France.10 As with other V. labrusca hybrids, winemakers prized 'Catawba' grapes for their versatility and hardiness.11 However, unlike many V. labrusca hybrids, 'Catawba' has less of a \\\"foxy\\\" or \\\"musky\\\" flavor, instead often described as a \\\"perfumey\\\" grape.12 The purple-lilac grapes, Adlum declared, were \\\"the most beautiful \u2026 to the eye, when they begin to ripen, that I know of.\\\"13 'Catawba' and the Ohio Wine Industry This \\\"most beautiful\\\" grape soon played a critical role in the expansion of the American wine industry and westward expansion more generally. Historians have long outlined the relationship between winemaking and imperialism. Erica Hannickel, in her book Empire of Vines, argues that grape culture in the nineteenth-century United States was distinctly expansionist, not only in terms of artistic and literary representations of the grapevine but also the physical space the vineyard occupied.14 As she argues, \\\"Americans believed if they could make good wine, everything else in terroir's web\\\"\u2014the \\\"totalizing ecology of the vineyard\u2014\\\" would be legitimated for their new country\u2014including its land, methods, farmers, and claim to international prestige.\\\"15 Here, Hannickel shows, \\\"grapes and the myth of terroir rooted the nation's imperial sense of itself\\\" during the period of westward expansionism. One man, Nicholas Longworth, argued in favor of Ohio's potential as an ideal location for grape-growing. Inspired by both the profit and potential improvements to the nation wrought by viticulture, Longworth began growing grapes as early as 1813 in his backyard.16 By 1823, Longworth was ready to set up a commercial endeavor. In that year, he began to cultivate his first vineyard, a four-acre plot.17 While Longworth began planting South African Cape grapes to make madeira, he soon transitioned to growing Vitis vinifera, which was already popular among American consumers. He went on to dedicate himself to V. vinifera grapes: in one letter to the editors of the Cincinnati Gazette, he writes that he worked for thirty years to cultivate V. vinifera grapes \\\"from all latitudes,\\\" to no success.18 Longworth's efforts reflect a broader preference for European wine grapes to the exclusion of wild American varieties, which stemmed from a belief in the superiority of V. vinifera. One nineteenth-century grape manual by viticulturist George Engelmann declared the European variety the \\\"only true\\\" wine grape and suggested that native American grapes were not worth cultivating due to their poor quality.19 However, Longworth soon found that Vitis vinifera was not easy to grow in the United States. In the thirty years Longworth attempted to cultivate European wine grapes, he claimed to \\\"have never found one worthy of cultivation in open air.\\\"20 This experience was not unique to Longworth but instead echoed throughout the nation. As one early American botanist claimed, despite the \\\"hundreds of thousands\\\" of European vines imported, they all failed due to the North American climate.21 Myriad horticultural manuals from the early nineteenth century confirmed that planting Vitis vinifera grapes in the United States was a herculean, if not impossible, task. winemaking and imperialism. Erica Hannickel, in her book Empire of Vines, argues that grape culture in the nineteenth-century United States was distinctly expansionist, not only in terms of artistic and literary representations of the grapevine but also the physical space the vineyard occupied.14 As she argues, \\\"Americans believed if they could make good wine, everything else in terroir's web\\\"\u2014the \\\"totalizing ecology of the vineyard\u2014\\\" would be legitimated for their new country\u2014including its land, methods, farmers, and claim to international prestige.\\\"15 Here, Hannickel shows, \\\"grapes and the myth of terroir rooted the nation's imperial sense of itself\\\" during the period of westward expansionism. One man, Nicholas Longworth, argued in favor of Ohio's potential as an ideal location for grape-growing. Inspired by both the profit and potential improvements to the nation wrought by viticulture, Longworth began growing grapes as early as 1813 in his backyard.16 By 1823, Longworth was ready to set up a commercial endeavor. In that year, he began to cultivate his first vineyard, a four-acre plot.17 While Longworth began planting South African Cape grapes to make madeira, he soon transitioned to growing Vitis vinifera, which was already popular among American consumers. He went on to dedicate himself to V. vinifera grapes: in one letter to the editors of the Cincinnati Gazette, he writes that he worked for thirty years to cultivate V. vinifera grapes \\\"from all latitudes,\\\" to no success.18 Longworth's efforts reflect a broader preference for European wine grapes to the exclusion of wild American varieties, which stemmed from a belief in the superiority of V. vinifera. One nineteenth-century grape manual by viticulturist George Engelmann declared the European variety the \\\"only true\\\" wine grape and suggested that native American grapes were not worth cultivating due to their poor quality.19 However, Longworth soon found that Vitis vinifera was not easy to grow in the United States. In the thirty years Longworth attempted to cultivate European wine grapes, he claimed to \\\"have never found one worthy of cultivation in open air.\\\"20 This experience was not unique to Longworth but instead echoed throughout the nation. As one early American botanist claimed, despite the \\\"hundreds of thousands\\\" of European vines imported, they all failed due to the North American climate.21 Myriad horticultural manuals from the early nineteenth century confirmed that planting Vitis vinifera grapes in the United States was a herculean, if not impossible, task. varieties are everywhere met with, springing up spontaneously in our woods and prairies, nature's own gift unaided by culture or by toil.\\\"25 For Prince, this underscores that the United States \\\"possess[es] not only all the advantages that France and other wine countries enjoy, from our having already introduced the choice varieties which those climes can boast, but this advantage is enhanced by the numerous varieties which our own country presents to us.\\\"26 Of the native grapes, Vitis labrusca emerged as the species of choice in Ohio for several reasons. For one, V. labrusca was indigenous to the American Northeast and could withstand dampness and cold\u2014 so much so that the grape allegedly failed when planted in the warmth of southern France.27 Further, \\\"the large size of the fruit, the vigor and productiveness of the vine, and its easy propagation from cuttings, made the varieties of this species preferable\\\" to others, according to one 1895 illustrated catalog on the American grapevine.28 These reasons transformed V. labrusca grapes into the grape of choice for early American viticulturists. As a result of a growing understanding of its qualities, 'Catawba' grape became a popular cultivar in the Northeast and Ohio regions. One 1871 article noted the widespread preference for the grape by both \\\"emigrants from the best wine districts of Europe, as well as those of American birth,\\\" based on \\\"careful analytical comparison with the best foreign wines, as well as upon taste, bouquet, &c.\\\"29 'Catawba' was also considered by some viticulturists to have a flavor superior to most indigenous grapes. In one letter to Adlum dated February 12th, 1827, a Dr. George Holcombe lauded the recipient for his role in popularizing 'Catawba': \\\"I congratulate you upon the success of your wines, particularly your Catawba, which is incomparably the best\u2014much the best sweet wine I have ever tasted.\\\"30 These reasons led viticulturists to declare that because of 'Catawba', America may someday become \\\"the vineyard of the west.\\\"31 Longworth, too, recognized the immense potential of the 'Catawba' in the Ohio context after Adlum sent him a sample in 1823. Longworth soon turned his focus to the 'Catawba' grape to establish the Cincinnati wine industry as the seat of wine production in the United States during the mid-nineteenth century. He wrote of 'Catawba', \\\"We have native grapes in most of our states, could a selection be made, which would leave us so little cause to regret that foreign grapes succeed so badly with us.\\\"32 And he began to note the advantages of the native grape over imported varieties, claiming in one profile that, unlike certain European wines, 'Catawba' does not \\\"run into the acetous fermentation, or become ropy.\\\"33 The few statistics we have from the 1830s suggest that Longworth's operation was initially small. In 1833, he owned nine vineyards which produced approximately 3,000 gallons of wine.34 By the 1840s, however, his scale of operations took off as he made a champagne-like \\\"sparkling Catawba\\\" which proved immensely popular throughout the nation and was immortalized in many poems. One Charles Hackay wrote an ode to the grape, ending with \\\"Catawba! Heart warmer! \/ Soul cheerer! life-zest! \/ Catawba, the nectar \/ And balm of the West.\\\" In 1850, Longworth produced 60,000 bottles of Catawba wine; that number rose to 75,000 by 1852.35 These bottles sold very quickly: one 1852 report noted that demand was \\\"much above the ability of Mr. L. to supply.\\\"36 Catawba had rapidly become one of the most popular\u2014and most quintessential\u2014 American wines. Historical Forces of Failure But even as 'Catawba' represented a successful departure from European wine grapes, most producers continued to employ European-style viticultural techniques. The 'Catawba' grape was raised in a highly regimented, monocrop culture typical of many European vineyards.37 Longworth also chose to use trellises to support his plants in the typical Rhineland style. Present-day viticulturists suggest a potential reason for this imitation of Rhine viticulture: whereas European vineyards in flat areas do not use trellises as the vines are able to support themselves, the younger vines of the United States could not stand alone, therefore requiring trellises like the grapes growing on the steep hills of the Rhine.38 Another potential reason was that most of Longworth's vineyards were run by German immigrants, many of whom came from the Rhine.39 Their experience growing grapes in Europe may have led them to adopt similar techniques upon arrival in Cincinnati, even though the topography of Ohio differed from that of their natal lands. This attempt to transform Cincinnati into the \\\"Rhineland of the West\\\" ultimately failed, in part due to this adherence to traditional European styles of viticulture. Because the 'Catawba' grape vines were planted so close to one another and monocropped, they were particularly susceptible to airborne fungal diseases and pests not initially found in Europe. The most destructive of these diseases originally found in America was black rot, which before the 1880s had only been observed in the Eastern United States. Black rot is a fungal disease that appears in the rainy season, as \\\"water activates the release of fungal spores.\\\"40 The spores cause the grapes to shrivel, making them unusable. By 1885, black rot was reported in France, most likely from the transfer of grape vines from the United States to Europe.41 However, this particular fungal disease had not been recorded on the European continent before that time. Because of the controlled, tight planting of the 'Catawba' grape, black rot deeply impacted the Ohio wine industry. According to a 1911 article on the wine industry in Ohio, \\\"From 1850 to 1880 it was difficult to get a bunch of any size and evenly ripened where it escaped the rot.\\\"42 Despite this problem, viticulturists like Longworth continued to praise 'Catawba' as the best of America's native grapes. It is unclear why Longworth remained so loyal to 'Catawba'; perhaps it was due to the popularity of sparkling 'Catawba' nationwide or fealty to the idea of producing wine from a grape native to America. By the 1860s, the tide began to turn against 'Catawba'. Viticulturist George Husmann, at a meeting of the Missouri Horticultural Society, allegedly tried to place 'Catawba' on the \\\"rejected list\\\" of vines, given its liability to various fungal diseases.43 And, just five years later, in 1865, one outlet lamented, \\\"We regret to learn that this variety of grape is nearly destroyed the present season by rot.\u2026 Every one having the Catawba in this vicinity, tell us a sad story about them in the present season.\\\"44 Here, then, we see a paradox of the early Ohio wine industry: even as producers like Longfellow opted for indigenous grapes rather than European vinifera, they stuck to traditional European cultivation mechanisms, leading to the industry's ultimate failure. Some wine producers recognized that the problems with 'Catawba' could be solved by departing from European viticultural methods. One \\\"E.P.C,\\\" writing in a popular Cincinnati newspaper in 1858, argued that the grape was sensitive to diseases only \\\"when not sufficiently aired and ventilated.\\\" The author underlines the failure of traditional planting techniques: \\\"That our old mode of close planting, heading-down, and crowding to the earth, with a view to get branches as near the ground as possible, is not calculated to remove this difficulty, is clear to all and self-evident.\\\"45 Anecdotal evidence suggests that he was correct: an 1888 contributor to a national agricultural journal notes that when he planted 'Catawba' with more room than normal, \\\"there has been no rot in twenty-eight years.\\\"46 Such alternatives demonstrate that 'Catawba''s proto-industrialized, regimented nature catalyzed the spread of black rot. However, vineyardists in Ohio, like Longworth, continued to plant 'Catawba' in the traditional Rhine style. It is unclear why this happened when national farming magazines and local papers printed information advising against monocropping 'Catawba' grapes so close together. Further, it is clear that alternative methods of grape-growing were present in other parts of the United States. Viticulturists in the Hudson Valley, for example, often practiced \\\"mixedfruit farming\\\" by planting red currants below the grapes and raspberries, strawberries, or a vegetable in between vineyard rows, with fruit trees at the end of every third row.47 The Ohio industry could have drawn upon these alternatives. Instead, perhaps due to attachment to European agricultural traditions, the monocultural cultivation of 'Catawba' pushed the Cincinnati wine industry towards its end. By the 1860s, the wine industry in Ohio folded in part due to fungal diseases.48 Its ravages, combined with the lack of laborers in vineyards due to the Civil War, as well as the death of Nicholas Longworth, meant that by 1870, grape-growing almost wholly vanished from Ohio.49 The Ohio wine industry\u2014and the 'Catawba' grape that undergirded it\u2014folded after just 50 years. Soon after, the behemoth we know today as the California wine industry took off in Ohio's stead. Other wine regions, like the Hudson Valley, also came into vogue as the cooler climate limited fungal pressures.50 The Catawba Wine Renaissance? But even as the Ohio wine industry failed, the 'Catawba' grape never totally disappeared, but is used in \\\"Pink Cats\\\" sweet wine as well as Kosher juice and wine.51 And today, Catawba wine is making a comeback. Skeleton Root, a popular Cincinnati winery, is \\\"focused on the revitalization of the local growing region,\\\" and makes delicious and unique Catawba wines that pay homage to Longworth's legacy. 52 As Skeleton Root winemaker Kate MacDonald noted, \\\"The West Coast is very forward about wine heritage, particularly in Sonoma where they have a lot of pre-prohibition wineries.\u2026 I became obsessed once I learned about Cincinnati's wine heritage.\\\" For MacDonald, using grapes like 'Catawba' is an opportunity \\\"to produce a wine that's uniquely American and uniquely Cincinnati.\\\"53 As MacDonald points out later in the interview, 'Catawba' offers more than a fascinating window into American agricultural and environmental history: it also offers a potential path forward in light of climate change. As the climate gets increasingly hotter and more unstable, fungal disease, which thrives on moisture, has the potential to impact grapes even more than normal. \\\"Instead of spraying [fungicides] like five times a year, which we would have done 15 or 20 years ago, we are having to spray 12 or 15 times a year,\\\" explains viticulturist and historian Stephen Casscles.54 \\\"I'm the one spraying these things, [so] I just would rather not use things that are highly toxic,\\\" Casscles says. \\\"Or, if I am, I'd rather use them three times a year rather than twelve.\\\"55 It is ironic that, despite the fact that 'Catawba' initially failed due to fungal disease, many winemakers are convinced that when grown in a more spaced out, polycultured manner, the grape can offer a potential antidote to frequent spraying. \\\"The labrusca varieties are marvels of biology\\\" says Phil Plummer, a winemaker at a Montezuma Winery in New York who works with 'Catawba' grapes. As he explains, V. labrusca grapes co-evolved with many of the fungal diseases and pests native to North America. \\\"If you can develop a hybrid,\\\" he notes, \\\"there are some of the characteristics and flavor and aroma of a V. vinifera, with some of the hardiness and disease resistance of the wild grapes that grow around here.\\\" This allows Plummer to successfully propagate grapes like 'Catawba' without \\\"breaking the bank or being out in the vineyard with the sprayer weekly.\\\"56 This is necessarily an important issue in environmental justice: Justine Belle Lambright, director of external business at the Kalch\u0113 Wine Cooperative in Vermont, points out, \\\"Although the majority of workers in a vineyard are Black and brown bodies, they only make up one percent of the ownership level.\\\" Hybrid grapes, which tend to require fewer chemical inputs, thus can improve health conditions for workers of color.57 While many historians have marked the Ohio wine industry as simply a failure, the long history of 'Catawba' suggests the vitality of the early American wine industry and a path forward for winemakers today. Hybrid grapes like 'Catawba', alongside other hybrids including 'Croton', 'Empire State', and more, allowed the industry to thrive for many decades. While 'Catawba' may have initially failed in the American wine industry, from its failure\u2014as well as its successes\u2014we might find new ways forward in cultivating crops during our current climate crisis. endnotes 1. Erica Hannickel, Empire of Vines: Wine Culture in America (Philadelphia: University of Pennsylvania Press, 2013), 105. 2. Bush, Son, and Meissner, Illustrated Descriptive Catalogue of American Grape Vines: A Grape Growers' Manual (St. Louis: R. P. Studley & Co., 1895), 99. 3. Henry Wadsworth Longfellow, \\\"Catawba Wine,\\\" in Birds of Passage (London: Routledge, 1878), 62. 4. Phil Plummer, Personal Interview, Phone, April 7, 2022; \\\"Catawba Grape Juice,\\\" Sweetwater Cellars, accessed June 20, 2022, https:\/\/sweetwatercellars.com\/catawbawhite. html; Julia Fine, \\\"Will Climate Change Help Hybrid Grapes Take Root in the US Wine Industry?,\\\" Civil Eats, June 16, 2022, https:\/\/civileats.com\/2022\/06\/16\/willclimate- change-help-hybrid-grapes-take-root-in-theus- wine-industry\/. 5. Richard White, Railroaded: The Transcontinentals and the Making of Modern America (New York: W. W. Norton & Company, 2011), 28. 6. John Frederic von Daacke, \\\"'Sparkling Catawba': Grape Growing and Wine Making in Cincinnati, 1800-1870\\\" (MA Dissertation, University of Cincinnati, 1964), 10 7. S. Downer, \\\"Native Grapes: Catawba Grape,\\\" The New England Farmer, and Horticultural Register 8, no. 29 (February 5, 1830): 226-27. 8. J. Stephen Casscles, Grapes of the Hudson Valley: And Other Cool Climate Regions of the United States and Canada (Coxsackie: Flint Mine Press, 2015), 70. 9. Ibid. 10. F. Huber et al., \\\"A View into American Grapevine History: Vitis Vinifera Cv. 'S\u00e9millon' Is an Ancestor of 'Catawba' and 'Concord,'\\\" VITIS - Journal of Grapevine Research 55, no. 2 (May 11, 2016): 53-56. 11. Casscles, Grapes of the Hudson Valley, 69. 12. Ibid., 69-70. 13. Downer, \\\"Native Grapes,\\\" 227. 1 4. Hannickel, Empire of Vines, 8. 15. Ibid., 12. 16. Pinney, A History of Wine in America, 157. 17. Ibid. 18. Robert Buchanan, The Culture of the Grape, and Wine-Making (Cincinnati: Moore, Anderson & Company, 1854), 106. 19. George Engelmann, The True Grape-Vines of the United States (St. Louis: R. P. Studley & Co., 1875), 2. 20. Buchanan, The Culture of the Grape, and Wine-Making, 106. 21. Engelmann, The True Grape-Vines of the United States, 2. 22. For more on these indigenous grapes, see: J. Stephen Casscles, Grapes of the Hudson Valley: And Other Cool Climate Regions of the United States and Canada (Coxsackie: Flint Mine Press, 2015). 23. Buchanan, The Culture of the Grape, and Wine-Making, 106. 24. William Robert Prince, A Treatise on the Vine: Embracing Its History from the Earliest Ages to the Present Day, with Descriptions of Above Two Hundred Foreign and Eighty American Varieties; Together with a Complete Dissertation on the Establishment, Culture, and Management of Vineyards \u2026 (New York: T. & J. Swords, 1830), v. 25. Ibid. 26. Ibid. 27. Paul Lukacs, American Vintage: The Rise of American Wine (New York: W.W. Norton & Company, 2005), 126. 28. Bush, Son, and Meissner, Illustrated Descriptive Catalogue of American Grape Vines: A Grape Growers' Manual, 27. 29. W. G, \\\"The Vineyard: The Catawba as a Wine Grape.\\\" Prairie Farmer (Chicago, February 11, 1871), 43. 30. John Adlum, A Memoir on the Cultivation of the Vine in America: And the Best Mode of Making Wine, 2nd ed. (Washington: Printed for the author, by William Greer, 1828), 150. 31. H. Shaw, \\\"The Catawba Grape,\\\" Prairie Farmer (Chicago, August 1849), 251. 32. Annual Report of the Ohio State Board of Agriculture: With an Abstract of the Proceedings of the County Agricultural Societies, to the General Assembly of Ohio \u2026 (Columbus: State Printers, 1860), 467. 33. Annual Report of the Commissioner of Patents (Washington: Wendell and Van Benthuysen, 1848), 463. 34. Pinney, A History of Wine in America, 161. 35. Ibid. 36. \\\"Grape Culture in the United States.,\\\" Valley Farmer (1849-1864) 4, no. 9 (September 1852): 314. 37. Hannickel, Empire of Vines, 105. 38. J. Stephen Casscles, Personal Interview, Phone, December 3, 2021. 39. Dann Woellert, Cincinnati Wine: An Effervescent History (Mount Pleasant: Arcadia Publishing, 2021). 40. Casscles, Grapes of the Hudson Valley, 30. 41. F. Lamson Scribner and Pierre Viala, Black Rot (Washington DC: Government Printing Office, 1888), 5. 42. Lowell Roudebush, \\\"The Catawba Grape,\\\" National Stockman and Farmer (Pittsburgh: Center for Research Libraries, 1911), 880-881. 43. \\\"HORTICULTURAL NOTES: Who Wants Pruning Well Done Father Abraham Apple Iron Hot-Bed Sashes An Inquiry The Catawba Not a First-Rate Grape Ellwanger & Barry's Catalogue Cherry Solons The Apple Tree Borer,\\\" Michigan Farmer (Lansing, February 25, 1860), 59. 44. \\\"Catawba Grapes,\\\" Colman's Rural World (St. Louis: July 1, 1865), 102. 45. E. P. C, \\\"More About the Grape,\\\" The Cincinnatus (Cincinnati: Center for Research Libraries, September 1, 1858), 412-414. 46. M. Clay, \\\"Grape-Rot,\\\" National Stockman and Farmer (Pittsburgh: Center for Research Libraries, September 13, 1888), 432-433. 47. Casscles, Grapes of the Hudson Valley, 10. 48. von Daacke, \\\"Grape-Growing and Wine-Making in Cincinnati\\\", 210. 49. Ibid. 50. See: Casscles, Grapes of the Hudson Valley. 51. Phil Plummer, Personal Interview, Phone, April 7, 2022; Dan Berger, \\\"The Feast of Unleavened Bread: Kosher Wine: A Buyer's Guide,\\\" Los Angeles Times, April 4, 1993. 52. \\\"Skeleton Root Winery,\\\" The Skeleton Root, urban winery and event space in Over the Rhine, Cincinnati, accessed July 1, 2022, http:\/\/www.skeletonroot.com\/about. 53. Karen Day, \\\"Interview: Winemaker Kate MacDonald, Skeleton Root Winery,\\\" Cool Hunting, January 8, 2018, https:\/\/coolhunting.com\/food-drink\/skeleton-root-winery\/. 54. Casscles, Personal Interview, Phone, December 3, 2021. 55. Julia Fine, \\\"Will Climate Change Help Hybrid Grapes Take Root in the US Wine Industry?,\\\" Civil Eats, June 16, 2022, https:\/\/civileats.com\/2022\/06\/16\/willclimate- change-help-hybrid-grapes-take-root-in-theus- wine-industry\/ 56. See ibid. 57. Ibid."},{"type":"arnoldia","title":"Trophic Cascade","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25801","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060af28.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"7","authors":"Dungy, Camille T.; Holten, Katie","start_page":"41","end_page":"41","article_content":"After the reintroduction of gray wolves to Yellowstone and, as anticipated, their culling of deer, trees grew beyond the deer stunt of the mid century. In their up reach songbirds nested, who scattered seed for underbrush, and in that cover warrened snowshoe hare. Weasel and water shrew returned, also vole, and came soon hawk and falcon, bald eagle, kestrel, and with them hawk shadow, falcon shadow. Eagle shade and kestrel shade haunted newly-berried runnels where mule deer no longer rummaged, cautious as they were, now, of being surprised by wolves. Berries brought bear, while undergrowth and willows, growing now right down to the river, brought beavers, who dam. Muskrats came to the dams, and tadpoles. Came, too, the night song of the fathers of tadpoles. With water striders, the dark gray American dipper bobbed in fresh pools of the river, and fish stayed, and the bear, who fished, also culled deer fawns and to their kill scraps came vulture and coyote, long gone in the region until now, and their scat scattered seed, and more trees, brush, and berries grew up along the river that had run straight and so flooded but thus dammed, compelled to meander, is less prone to overrun. Don't you tell me this is not the same as my story. All this life born from one hungry animal, this whole, new landscape, the course of the river changed, I know this. I reintroduced myself to myself, this time a mother. After which, nothing was ever the same. camille t. dungy is the author of the essay collection Guidebook to Relative Strangers: Journeys into Race, Motherhood, and History, and four collections of poetry, most recently Trophic Cascade. She has edited three anthologies, including Black Nature: Four Centuries of African American Nature Poetry. Her honors include the 2021 Academy of American Poets Fellowship, a Guggenheim Fellowship, an American Book Award, and fellowships from the NEA. She is a University Distinguished Professor at Colorado State University. katie holten is an artist and activist. In 2003, she represented Ireland at the Venice Biennale. Exhibiting internationally, she is the recipient of numerous grants and fellowships, including a Fulbright and a MacDowell Fellowship. In her book The Language of Trees (Tin House, April 2023), Holten uses a \\\"tree alphabet\\\" of her own design to interpret the work of more than fifty contributors, including Dungy's \\\"Trophic Cascade.\\\""},{"type":"arnoldia","title":"The Roots of Rejuvenation","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25802","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060af6c.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"6","authors":"Del Tredici, Peter","start_page":"30","end_page":"40","article_content":"I've been studying how trees can rejuvenate themselves through sprouting since 1989, when I traveled to Tianmu Mountain in eastern China to study a wild population of Ginkgo biloba and discovered that many of the ancient specimens growing there had produced secondary trunks in response to storm damage, logging, or landslides. Twelve years later, I published a review article describing the morphological mechanisms that temperate trees have developed for generating basal shoots following traumatic disturbance, from root suckers and branch layers to stump sprouts and lignotubers. Essentially, sprouting is a form of clonal growth that not only allows woody plants to recover from damage but also to circumvent the ravages of aging, especially when they are able to produce new, adventitious roots to support their new shoots. In botanical sciences, the word adventitious is traditionally used to describe a plant structure that is somehow \\\"out of place,\\\" such as a shoot produced by a root, or a root produced by a stem. Within the field of plant propagation, the term is used to describe roots produced by detached stem cuttings treated with auxins under greenhouse conditions. Intact trees growing in natural habitats also produce adventitious roots from their stems, either as a genetically programmed part of their normal development\u2014such as the free-hanging aerial roots produced by tropical trees like strangler figs and mangroves\u2014or induced by changing environmental conditions such as partial uprooting or burial with silt following flooding. Regardless of whether they are programmed or induced, adventitious roots always develop from cells that neighbor vascular tissues located just below the bark (Bellini et al. 2014). Types of Adventitious Roots My research has shown that the trunks and branches of intact temperate trees can produce five different kinds of adventitious roots, the most common of which are those that develop on low-hanging lateral branches that come in contact with soil\u2014a process known as layering. The phenomenon has been documented in many different species of trees and is particularly common among conifers, both wild and cultivated. In many old estates, trees grown as widely spaced specimens often retained their lower horizontal branches that, under the influence of age and gravity, ended up resting on the ground and taking root. Once they take root, the branches change from a horizontal to a vertical orientation and, over time, can form a ring of new trees surrounding the original parent trunk. There is an extensive literature on layering, which David Orwig of the Harvard Forest and I reviewed in 2017 in our article about this behavior in the eastern hemlock, Tsuga canadensis. Genera with species that commonly form layers include Chamaecyparis, Fagus, Picea, Platanus, and Thuja. Lateral branches high up in the tree canopy also produce adventitious roots when they are covered with moisture-trapping epiphytes. As ecologist Nalini Nadkarni has shown, such \\\"canopy roots\\\" play an important role in nutrient absorption and cycling in tropical rain forests and, as described by Dietrich Hertel, to a more limited extent in temperate forests among such genera as Acer, Alnus, Fagus, and Populus. In wetland habitats, the partially uprooted trunks of trees lying on the ground often produce adventitious roots that allow the prostrate stem to generate new vertical shoots, a phenomenon known as trunk layering. In a similar vein, the vertical trunks of trees that have been partially covered with soil, silt, or water after flooding often produce adventitious roots that allow them to adjust to the new conditions, as seen in species in the genera Alnus, Larix, Salix, Sequoia, and Taxodium. Many trees produce swollen root collars with the capacity to generate adventitious roots as well as secondary shoots following some form of traumatic disturbance. In some long-lived species, these basal swellings\u2014technically known as lignotubers\u2014are a genetically programmed part of normal seedling development. In others, the they are induced by repeated coppicing or some other type of environmental disturbance. Examples of the former include species in the genera Ginkgo, Eucalyptus, Olea, Sequoia, and Tilia while the latter include species in the genera Betula, Castanea, Morus, Platanus, and Quercus. As stated above, my interest in lignotuberproducing trees began with my studies of ginkgo trees in eastern China, where I observed that many of the ancient specimens growing on Tianmu Mountain had produced secondary stems in response to damage to their primary trunks. My research on lignotubers continued in California where the coast redwood produces massive underground lignotubers that, following logging, can generate \\\"fairy rings\\\" of new trunks that can extend the tree's lifespan for centuries (Del Tredici, 1992, 1998, 1999; Del Tredici et al. 1992). The fifth and final type of adventitious root\u2014and the focus of the remainder of this article\u2014are those that emerge from the woundwood that trees produce when their trunk is damaged and its xylem exposed. In response to this damage, trees generate undifferentiated callus tissue that gives rise to a new layer of vascular cambium that produces the woundwood that will eventually cover over the injury (Stobbe et al. 2002). During the healing process, woundwood typically grows inward from the edges of the damage and continues growing until its margins come together to seal off the exposed wood. If the wood to the interior of this callus tissue is firm, this is usually the end of the process. If the wood is rotten, however, it offers little resistance to the expanding woundwood which rolls inward on itself and continues growing inside the trunk. In response to the moist, dark conditions within the trunk, the vascular cambium of the woundwood can initiate adventitious roots that grow into its own rotten core. If the rot extends through the trunk down to the ground, these roots have the capacity to generate a new subterranean root system and develop into stout columns that can provide extra support for the hollow trunk. While many angiosperm trees have been reported to form internal trunk roots, it is a relatively uncommon phenomenon that occurs mainly in old, open-grown specimens that have experienced extensive branch loss. Internal Trunk Roots in the Literature Descriptions of internal trunk roots are rare in the botanical and horticultural literature. The earliest mentions of the phenomenon that I could locate were published in the British periodical Gardeners' Chronicle and Agricultural Gazette between 1853 and 1877. William Booth wrote the first article and reported finding them inside the trunks of sweet chestnut (Castania sativa) and Cornish elm (Ulmus minor 'Stricta'). In both cases, the roots originated from woundwood and reached down to the ground. A second article, authored by \\\"Vigilax,\\\" appeared a week later and described the same phenomenon in a specimen of Laburnum. The third reference to internal trunk roots is by Moggridge in 1870, who described four old pollarded pedunculate oaks (Quercus robur) growing in Richmond Park in Surrey, England, that had produced large internal, adventitious roots, two of which he illustrated. The final Gardeners' Chronicle article on the subject is from 1877 by W. G. S., who described a large internal trunk root in an ancient specimen of yew (Taxus baccata). In the modern scientific literature, the first reference to internal trunk roots is from 1908 by O. M. Ball, who reported their existence in an old specimen of the umbrella chinaberry, Melia azedarach 'Umbraculifera'. He called the process \\\"self-eating\\\" and described how \\\"roots descend through the decaying materials and often, upon reaching the harder, less decayed wood of the lower part of the stump, turn sharply back and grow upward even to the point of origin.\\\" He published a photograph of the specimen that shows the roots originating from woundwood. Such was the extent of the literature until 1954, when Czech botanist Jen Jenik published a short article describing internal trunk roots in European beech, Fagus sylvatica. In 1975, another Czech scientist, Jarmila Kubikov\u00e1, published an article that not only described their morphology in detail but also the amazing ecosystem inside the trunks of rotten linden trees (Tilia). To this day, Kubikov\u00e1's summary of his research remains the one of the best descriptions of internal trunk roots. Noting that the phenomenon is \\\"little known,\\\" he describes the \\\"specific ecosystem\\\" found within old trees, which includes \\\"saprophytic fungi and bacteria, together with numerous protozoa, snails and various groups of insects.\\\" Internal roots only develop when the living material of the cavity \\\"reaches the living peripheral tissues of the trunk whose meristems form the healing callus\\\" from which root primordia eventually develop. Kubikov\u00e1 described their significance in older trees as \\\"important regeneration phenomena which extend the number of absorption rootlets, improve the transport of water and nutrients, and thus enable the growth of new branches and photosynthetic apparatus. At the same time, these roots function as supplementary armature increasing the resistance of a tree to damage by wind and snow. Thus the ageing process can be retarded and the life span of the tree prolonged.\\\" Three years later, in 1978, Dickenson and Tanner reported the occurrence of roots inside the hollow trunks of several species of Jamaican trees that were cut down in an experimental logging operation. Ten of the thirty-nine cut trees were hollow and \\\"many\\\" of them contained roots. They examined four trees in detail and found that in only one case the roots were produced by the tree itself. In the other three individuals, the internal trunk roots were produced by epiphytes attached to the branches of the tree or by nearby fig trees (Ficus sp.) whose roots grew a meter and a half upwards into the rotten trunk from the mineral soil below. The authors concluded that they had found \\\"examples of a possible advantage gained from having a hollow trunk and examples of a possible disadvantage.\\\" Their observations are important because they remind us that the nutrients inside a tree's rotten core are fair game for any plant than can reach them from above or below. Remarkably, it was not until 1992 that internal trunk roots were given a proper scientific name by two Chinese scientists, Liu Qijing and Wang Zhan. Working on the windswept slopes of Changbai Mountain in northeast China\u2014which I visited in 1997\u2014they described their occurrence in numerous old specimens of Betula ermanii and called them \\\"endocaulous roots\\\" in the English abstract of their article (which was written in Chinese). The study site was between 1700 and 2000 meters elevation with abundant annual precipitation (967-1400 mm) and fog. Because of strong winds and heavy snow, many of the birch trees were growing diagonally rather than vertically as they did at lower elevations. The authors looked at a large number of specimens and determined that endocaulous roots were common in trees around two hundred years old and 20 cm DBH (diameter at breast height, 4.3 feet above ground), and that these roots were able to grow down into the soil because their trunks had been hollowed out by extensive heart-rot. In younger trees, those less than 10 cm DBH, endocaulous roots were not common, and rarely reached the soil as there was much less heart-rot than in larger trees. In the moist forests on the north side of the mountain, trees greater than three hundred years old commonly developed endocaulous roots that reached the soil\u2014up to five or more per trunk\u2014some of which were more than 15 cm across and over one hundred years old. The extreme weather on Changbai Mountain caused serious damage to the trunks and branches of older trees which, in turn, promoted the development of heart-rot on the inside and woundwood on the outside. When the woundwood came into contact with the soft, moist heart-rot, it generated adventitious roots that grew into it and eventually made their way down into the soil. Liu and Wang made the additional observation that, \\\"Such rotten trees will be blown down or broken during wind storms so that endocaulous roots become visible. However, these broken trees will not likely die. Rather, their vigor increases because of the rapid development of endocaulous roots following the disturbance\\\". Unaware of the work of Liu and Wang, Jenik proposed calling the internal trunk roots, \\\"endocormic roots\\\" in 1994 based on the work of Kubikov\u00e1 from 1975 as well as his own earlier research from 1954. Remarkably, literature and internet searches of both terms show that neither of them have had a significant impact on the botanical or horticultural literature, a situation I intend to remedy with this article. The prefix caul- comes from the Latin caulis meaning stem or stalk, and corm- from the Greek kormos meaning tree trunk. While both terms are appropriate, I prefer endocormic for three reasons: 1) it was proposed by Jenik, who published an early description of the phenomenon in 1954; 2) it is derived from the same root as the widely used term, epicormic shoots, which describes new branches that sprout from an old trunk; and 3) it suggests the use of a new term, epicormic roots, to describe those that are produced by the trunk or branches of a tree when they come in contact with the soil. Observations on Endocormic Roots I observed my first endocormic roots in 1986, in a storm-damaged red oak, Quercus rubra, at the Arnold Arboretum that was being cut up for removal. Midway through the process, a member of the grounds crew noticed an unusual structure inside the trunk and called me over to look at it. What I saw amazed me\u2014the woundwood that had initially covered an old branch scar had turned inward and continued growing inside the trunk where it formed a large mushroom-shaped structure that had proliferated adventitious roots. I have been on the lookout for endocormic roots ever since, and have observed them in various gardens and parks in a number of old trees growing as isolated specimens of various types, including: Acer platanoides, Cladrastis kentuckea, Cornus controversa, Fagus sylvatica, Ginkgo biloba, Gymnocladus dioicus, Liriodendron tulipifera, Malus sp., Morus alba, Prunus \u00a9 yedoensis, Tilia americana, and Tilia \u00a9 vulgaris. Several authors have published illustrations of endocormic roots, including Oldeman (1990) in Fagus grandifolia; Mattheck (1991) in Fagus sylvatica; Thomas (2000) in Ulmus \u00a9 hollandica; Fay (2002) in Carpinus betulus, and Bragg (2018) in Acer rubrum. Writing in Arnoldia in 2012, Tony Aiello described an unusual specimen of Prunus subhirtella 'Pendula' at the Morris Arboretum in Philadelphia, in which a large endocormic root morphed into the stem of a stand-alone tree. Taken together, these reports indicate that the woundwood of most angiosperm trees has the capacity to generate endocormic roots when it comes into contact with rotten heartwood. For some unknown reason, internal trunk roots seem to be less common among gymnosperms. In New England, I have observed endocormic roots on several species of wild-growing trees, including sugar maple (Acer saccharum), red oak (Quercus rubra), and gray birch (Betula populifolia). Mostly the roots were confined to the cavities filled with rotten heartwood, but in a few cases the roots extended all the way down into the ground and developed into thick columns that helped support the hollow trunk. It seems likely that upon reaching the soil, these roots produce tension wood that causes them to contract and thicken and, over time, provide an added measure of structural support for the hollow trunk. Assuming this is the case, these column roots are probably behaving similarly to the aerial roots of Ficus that produce reaction wood that contracts when they reach the ground (Gill & Tomlinson, 1975). On August 4, 2020, I had the rare opportunity to observe multiple cases of endocormic root formation after tropical storm Isaias passed near the town of Cornwall, Connecticut where I was spending the summer, and seriously damaged many of the trees growing along the roadways. While Isaias caused lots of problems for the people who lived there\u2014power was out for ten days\u2014it offered the opportunity to observe the condition of the trees that brought the power lines down, including several old specimens of sugar maple that had been planted over a century ago for syrup production. Most of these open-grown sugar maples that lined many of the roads possessed whorls of large, upright lateral branches about three meters up on their trunks, which showed an increasing tendency to brake off in storms as they got bigger. To my surprise, many of the large laterals that broke off the old sugar maples during Isaias revealed \\\"humus\\\" inside the trunks that was permeated with endocormic roots that had originated from woundwood produced in response to earlier limb loss. My observations on the morphology of opengrown sugar maples echoed the findings of Ranius and his colleagues who described hollow formation in the trunks of pedunculate oak (Quercus robur) growing in southwest Sweden in 2009. The authors found that among trees less than one hundred years old, fewer than one percent contained hollows, while fifty percent of trees between two and three hundred years old had hollows. They also observed that hollows formed at an earlier age in faster growing trees located in open pastures than in slower growing trees located in forests. The authors attributed this to the fact that most hollows were generated when large branches broke off and that such large branches formed sooner on open-grown trees than on forest-grown trees of the same age. Management of Ancient Trees There are long-standing questions in both the ecological and horticultural literature as to why so many trees have hollow cores and whether it compromises their stability. In 1976, Daniel Janzen was far ahead of his time in proposing that \\\"the rotten hollow core is often an adaptive trait, selected for as a mechanism of nitrogen and mineral trapping. A rotten core is a site of animal nests, animal defecation, and microbial metabolism that should result in a steady fertilization of the soil under the base of the tree.\\\" Graeme Ruxton in 2014 proposed an economic rather than a nutritional answer to the question, asserting \\\"the central wood of trees is allowed to decay because the costs of chemically defending it are not justified by the small reduction in structural stability that is likely to occur.\\\" To support his theory, he cited arboricultural research showing that if the radius of the inner hollow region of the trunk is less than seventy percent of the total radius, there is little cost to the tree in terms of reduced structural stability (Mattheck, 1994; Fink, 2009). Over the past twenty years, a considerable body of research has developed on the horticultural management of ancient and veteran trees. In particular, there is an extensive literature focusing on pruning techniques that promote the long-term survival of such trees while simultaneously creating habitat for the hollow-inhabiting (saproxylic) organisms that are dependent on them (Fay, 2002, Read et al., 2010, Gough et al., 2014, Fay & de Berker, 2016, Hirons & Thomas 2018, Bengtsson et al., 2021). One particularly interesting study by Pavel Sebek and his colleagues from 2013 looked at the incidence of hollows in pollard versus non-pollard white willow, Salix alba, in the Czech Republic. The authors sampled 1,126 trees across in four different areas and observed hollows in 83% of the pollarded trees and only 34% of the unpollarded trees. For trees of 50 cm DBH, they found that \\\"the probability of hollow occurrence was 75% in pollards, but only 30% in non-pollards.\\\" They concluded that actively managing trees via pollarding could speed up the creation of habitats for saproxylic organisms and contribute significantly to the conservation of rare, hollowdependent fauna. While the authors make no mention of endocormic roots, their research suggests that pollarding might well induce their development by virtue of the fact that it typically stimulates extensive woundwood formation. This modern approach to promoting the development of tree hollows for conservation purposes is a complete reversal of the once common practice of filling them with cement. In addition to promoting the development of tree hollows, pollarding is known to increase the lifespan of many trees. As Oliver Rackham noted in 1990, \\\"Trees whose function is not timber\u2014pollards and coppice stools\u2014may live much longer than timber trees. The cutting process prolongs their lives, and they go on doing their job of producing useful crops of poles despite old age or decay.\\\" Taking this into account, the repurposing of pollarding for conservation rather than production purposes is a \\\"winwin\\\" management strategy for both the trees and the organisms that live within their rotten hearts. In conclusion, the formation of endocormic roots by old trees is a manifestation of the senescent phase of their growth. Writing in 2013, Howard Thomas vividly describes senescence as a state in which the boundary between life and death is often blurred, as a tree seeks \\\"to control its own viability and integrity\\\" while confronting the \\\"thermodynamically unavoidable\\\" eventuality of death. From this perspective, a tree's ability to transform rotten heartwood into living tissue is an incredible example of how trees can navigate the ambiguity of their mortality by generating adventitious roots. acknowledgements The author expresses thanks to Jianhua Li of Hope College for help with Chinese translations and Michael Dosmann of the Arnold Arboretum for his support and thoughtful review of the manuscript. peter del tredici worked in a variety of capacities at the Arnold Arboretum for 35 years and taught at the Harvard Graduate School of Design and at MIT for over 20 years. His recent work is focused on urban ecology and climate change and he is the author of Wild Urban Plants of the Northeast: A Field Guide (2nd ed. 2020). An expanded version of this article was published in June 2022 in Arboricultural Journal. roots references Aiello, A. S. (2012). Japanese flowering cherries\u2014a 100-year-long love affair. Arnoldia, 69(4), 2-14. Ball. O. M. (1908). Formation of adventitious roots in the umbrella China tree. Botanical Gazette, 46(4), 303-304. Bellini, C., D. I. Pacurur & I. Perrone. (2014). Adventitious roots and lateral roots: similarities and differences. Annual Review of Plant Biology, 64, 17.1-17.28. Bengtsson, V., C. P. Wheater, H. Read & R. Harris. (2021). Responses of oak pollards to pruning, Arboricultural Journal, 43(3), 156-170. Booth, W. B. (1853). Curious instances of the formation of roots. Gardeners' Chronicle & Agricultural Gazette, 13 (January 1), 4. Bragg, D. C. (2018). A woody chamber of secrets. Frontiers in Ecology & Environment, 16(10), 598. Del Tredici, P., & D. A. Orwig. (2017). Layering and rejuvenation in Tsuga canadensis (Pinaceae) on Wachusett Mountain, Massachusetts. Rhodora, 119, 16-32. Del Tredici, P., H. Ling, and G. Yang. 1992. The Ginkgos of Tian Mu Shan. Conservation Biology 6(2): 202-209. Del Tredici, P. (1992). Natural regeneration of Ginkgo biloba from downward growing cotyledonary buds (basal chichi) American Journal of Botany, 79, 522-530. Del Tredici, P. (1998). Lignotubers in Sequoia sempervirens: development and ecological significance. Madrono, 45, 255-260. Del Tredici, P. (1999). Redwood burls: immortality underground. Arnoldia, 59(3), 14-22. Del Tredici, P. (2001). Sprouting in temperate trees: a morphological and ecological review. Botanical Reviews, 67(2), 121-140. Del Tredici, P. 2022. Endocormic roots: transforming death into life. Arboricultural Journal, DOI: 10.1080\/03071375.2022.2085943 Dickinson, T. A., & E. V. J. Tanner. (1978). Exploitation of hollow trunks by tropical trees. Biotropica, 10(3), 231-233. Fay, N. (2002). Environmental arboriculture, tree ecology and veteran tree management. Arboriucultural Journal, 26, 213-238. Fay, N., & N. de Berker. (2016). Ancient trees and their value. In K. Witko\u015b-Gnach & P. Tyszko-Ghenienlowiec (Eds.), Trees\u2014a lifespan approach (pp. 103-131). Wroclaw: Fundacja EkoRozwoju. Fink, S. (2009). Hazard tree identification by visual assessment (VIA): scientifically solid and practically approved. Arboricultural Journal, 32, 139-155. Gill, A. M., & P. B. Tomlinson. (1975). Aerial roots: an array of forms and functions. In J. G. Torrey and D. T. Clarkson (Eds.), The development and function of roots (pp. 237-260). New York, NY: Academic Press. Gough, L. A., T. Birkemoe & A. Syerdrup-Thygeson. (2012). Reactive forest management can also be proactive for wood-living beetles in hollow oak trees. Biological Conservation, 180, 75-83. Hertel, D. (2011). Tree roots in canopy soils of old European beech trees\u2014an ecological reassessment of a forgotten phenomenon, Pedobiologia, 54, 119-125. Hirons, A. D., & P. A. Thomas. (2018). Applied tree biology. Oxford, UK: John Wiley & Sons. Janzen, D. H. (1976). Why tropical trees have rotten cores, Biotropica, 8, 110. Jenik, J. (1954). Ko\u0159enov\u00fd syst\u00e9m vo kmeni buku [The root system in the beech trunk]. Vesmir, 33, 350-351 [In Czech]. Jenik, J. (1994). Clonal growth in woody plants: a review. Folia Geobotanica & Phytotaxonomica, Praha, 29, 291-306. Kubikova, J. (1975). Adventivn\u00ed ko\u0159enov\u00fd syst\u00e9m v dutin\u00e1ch star\u00fdch strom\u016f a jeho v\u00fdznam [Adventitious root systems in the cavities of old trees and its significance]. Preslia, 47, 331-334 [In Czech with English summary]. Liu, Q., & Z. Wang. (1992). Root system inside heart-rot stem of Betula ermanii. Research Forest Ecosystems, 6, 68-71 [In Chinese with English summary]. Mattheck, C. (1991). Trees: The mechanical design. Berlin: Springer-Verlag. Mattheck, C., & H. Breloer. (1994). The body language of trees. London: The Stationary Office. Moggridge, J. T. (1870). Pollard oaks. Gardeners' Chronicle & Agricultural Gazette, 30(Sept. 17), 1248-49. Nadkarni, N. M. (1994). Factors affecting the initiation and growth of above ground adventitious roots in a tropical cloud forest tree: an experimental approach. Oecologia, 100, 94-97. Nadkarni, N. M. (1981). Canopy roots: convergent evolution in rainforest nutrient cycles. Science, 214, 1023-1024. Oldeman, R. A. A. (1990). Forests: elements of silvology. Berlin: Springer-Verlag. Rackham, O. (1990). Trees & woodland in the British landscape (revised ed.) London: Phoenix Giant. Ranius, T., M. Niklasson & N. Berg. (2009). Development of tree hollows in pedunculate oak (Quercus robur). Forest Ecology & Management, 251, 303-310. Read, H. J., C. P. Wheater, V. Forbes & J. Young. (2010). The current status of ancient pollard beech trees at Burnham Beeches and evaluation of recent restoration techniques. Quarterly Journal of Forestry, 104(2), 109-120. Ruxton, G. D. (2014). Why are so many trees hollow? Biology Letters, 10: 20140555. Sebek, P., J. Altman, M. Platek & L. Cizek. (2013). Is active management the key to the conservation of saproxylic biodiversity? Pollarding promotes the formation of tree hollows. PLOS One, 8(3), e6045. Stobbe, H., U. Schmitt, D. Eckstein & D. Dujesiefken. (2002). Developmental stages of fine structure of surface callus formed after debarking of living lime trees. Annals of Botany, 89, 773-782. Thomas, P. (2000). Trees: their natural history. Cambridge: Cambridge Univ. Press. Thomas, H. (2013). Senescence, aging and death of the whole plant. New Phytologist, 197, 696-711. Vigilax. (1853). Curious formation of roots above ground. Gardeners' Chronicle & Agricultural Gazette, 13(Jan. 8), 21. W. G. S. (1877). Ancient resuscitated yew at Bettws Newydd. Gardeners' Chronicle, 7(Feb. 17), 215."},{"type":"arnoldia","title":"Oak and Pine in Life and Death","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25803","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060b36f.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"5","authors":"Carlson, Robin Lee","start_page":"16","end_page":"29","article_content":"In The Cold Canyon Fire Journals: Green Shoots and Silver Linings in the Ashes, artist and naturalist Robin Lee Carlson shares the six years she spent learning from the wildfires that burned through Stebbins Cold Canyon Reserve near Davis, California twice in five years. This excerpt, drawn from the chapter \\\"The Relativity of Time,\\\" explores the tree-entangled life that returns in abundance after fire\u2014some of which emerges rapidly, and some of which takes longer to make itself known. \\\"It is tempting to immediately tally the living and the dead,\\\" notes Carlson. \\\"But trees exist on a very different time scale, and their lives and deaths caution against hasty conclusions.\\\" \\\"Look at this burned bay laurel. Do you see the way those branches are all pointing the same direction?\\\" Dead tree branches just over my head etch lines into the electric-blue sky, and I am transfixed. From the thicker supporting branches to the finest twigs, they all swirl in the same direction, performing a ghostly dance choreographed by the Wragg Fire's fierce, hot wind.* Even though the air around me is completely still, I can easily envision the wind whipping around these trees as the fire blazed and they were frozen in this shape. It has been a year-and-a-half since the fire, and a participant in a field sketching class I am leading at Cold Canyon has just pointed out these shapes in the trees to me. Miriam Morrill is a biologist and a specialist in fire and communications at the Bureau of Land Management (BLM). She is also an artist, and I see that she has been making notes and drawings in her sketchbook that capture some of the signs of fire behavior still legible in the landscape. This is not a language that I know, and we are excited to compare notes. She explains that she sees more trees and tree skeletons on Blue Ridge than she would expect had the fire advanced up that slope from the bottom of the canyon. She surmises that the fire instead came from the west, over the top of the ridge, and backed down into the canyon, burning less intensely moving downslope than it would have had it been charging upslope. I tell her that this is what I know to have been the case, based on Jeffrey Clary's descriptions of the fire's progress. Miriam says that she also notices that the branches of the leafless trees where we are standing near the creek are all pointing in different directions, as if different eddies of wind were whirling around each one. From this, Miriam concludes that the fire became more intense at the bottom of the canyon before racing up the slope on the other side. As the Wragg Fire paused here, the intensity of the fire created its own weather. Strong winds gusted and eddied among the folds and pockets of the hills. Where the fire did not fully burn the vegetation, its heat dried out the branches. Losing their moisture made them permanently inflexible, pointing in the direction of the last wind they would ever feel. This phenomenon is termed foliage freeze. Time stopped, arrested, and I am held there too, caught in this still, quiet day but also in the midst of the roaring frenzy of fire and wind of late July 2015. This could be the image of violence preserved, jarring me out of time on a peaceful day. I might see this as the life force of these trees sucked right out of them by the wind, with their macabre skeletons still here to remind me. If fire is only a catastrophe from which this ecosystem must recover, the brutality of that image is appropriate. But if fire's place in the ecosystem is subtler, then those frozen branches might equally be seen as a reminder of the continuity that underlies the fire, tying the seemingly singular event to the past and future of the landscape. Disturbance has a past and a future and blends more easily into both than it would seem. And now, as I stand here in the present, the frozen branches are important clues to the behavior of the fire. Fire leaves signs behind on the land that are often fleeting\u2014 erased by wind and rain and footsteps\u2014so it feels like a race to find and read them before they disappear. Patterns of ash and char, curled leaves, fallen tree trunks, and grass stems\u2014all are quickly obscured by the movement of animals, the fall of rain and rush of wind, and the growth of new vegetation. These twisted branches, though, remain. The fire's passage is frozen in the trees' bodies, and the trees' writhing limbs marking this path are emblazoned on my mind. Oaks: Anchors in time Growing up in California's Central Valley ensured that water worries indelibly marked my subconscious. I am a child of brown summers and green winters, of regular drought and water scarcity. I was a toddler during the 1976-77 drought and have vivid memories of my mother explaining to me why we couldn't flush the toilet every time we used it. Even as a child, I approached each winter with concern: Will there be enough rain this year? Will the rivers dry up? I find nothing more reassuring than rain and nothing more terrifying than the endless cloudless summer skies. As I got older, I began to understand the ways life here has adapted to make the most of the rain we do receive. I am reassured by the hardy species that get by on little water and conserve their moisture in ingenious ways, like the blue oaks standing tall in the woodlands and savannas that ring the Central Valley. These are important members of Cold Canyon habitats, the trees that Jeffrey was concerned about after the Wragg Fire. Blue oaks are of middling size for oaks, with the wildly crooking branches common among oaks that make their limbs look as though they are dancing ecstatically. Their leaves are compact, with only wavy margins, not the deep lobes of some of the other oak species. The waxiness of their leaves helps with moisture retention in their oftenarid habitats. Staunchly Californian, blue oaks are endemic here\u2014found nowhere else in the world. They live at lower elevations in the Coast Ranges and the Sierra Nevada, as well as on the southern end of the Cascade and Klamath Mountains. A few grow in the Central Valley itself, but they mostly prefer to be up in the hills. Though I live in the flatlands of the valley, I am glad to be close to the hills and mountains, and it is to these foothill habitats that I turn for guidance in how to live in an often-parched land. Of all the deciduous oaks in California, blue oaks are the most able to withstand drought. Their taproot\u2014their main central root\u2014can grow deep into the ground to find the water table, to depths of eighty feet, though they can also focus their energy on growing shallower roots if water is available higher in the soil. They need soil that is relatively dry and well drained, and are found on soils that are poorer in nitrogen, phosphorous, organic matter, and other nutrients than the soils that support other California oaks. On these soils, they cultivate communities of other hardy species by holding the earth in place against erosion and using their taproots to bring water to the surface, incidentally making it available to other species as well. Before introduced annual species came to dominate California grasslands and savannas, blue oaks grew alongside bunchgrasses such as blue wildrye and purple needlegrass, similarly water-conserving species. The oaks and grasses mutually supported one another, sharing resources via fungal mycelial networks. Now, introduced annuals such as redstem filaree, cheatgrass, and wild oat have largely replaced the bunchgrasses. The introduced species compete with young oaks for space, water, and light, just as they outcompeted the native bunchgrasses for these resources. As in the rest of their range, blue oaks in Cold Canyon shelter quite a few introduced annual species, but they are also home and sustenance to many other creatures. Nuttall's woodpeckers rely on them for foraging and nesting. Dusky-footed woodrats build at their feet and eat their acorns. Fallen branches are important gathering places for western fence lizards. Oak titmice\u2014birds the soft mousy brown of their namesakes\u2014defend year-round territories in their canopies. It is easy to see the wildlife that depends on the oaks, but equally important to understand the less visible forces at work. Blue oaks are critical to the quality of the soil around them, increasing its nutrient content. This is thanks to the fall of leaf litter, which increases the nitrogen in the soil. It is also because trees capture moisture from the air that otherwise would have evaporated. Instead, the water droplets adhere to the leaves, eventually aggregate into larger drops and then fall to the ground. The drops bring along important nutrients\u2014 potassium, phosphorous, and magnesium, for example\u2014that were present in the air and are concentrated when the drops form. There are three kinds of oaks growing in Cold Canyon, two of which are trees\u2014blue oak and interior live oak\u2014and scrub oak, which takes a shrubbier form. Interior live oaks tend to be found in the bottoms of the canyons, as they prefer the wetter areas in the riparian zones, and these are often too wet for blue oaks. Blue oaks generally grow on the hillsides. The distinctions are not absolute: there are certainly blue oaks growing in the canyons and live oaks on the hills, but the general pattern holds. Over the five years of my visits, when I look closely at oaks, I try to decipher the puzzle of their health and survival. In a landscape of rapidly sprouting annual plants and fast-growing shrubs, trees are not as easy to assess. There are no immediate answers to how the oaks at Cold Canyon are faring. In the areas closest to the creek, some oaks did not burn at all and are doing fine\u2014these are mostly interior live oaks. A little further from the creek, there are more interior live oaks, and some of these burned. These that burned are sprouting, strong shoots growing up from their bases, from their root crowns. The brand-new leaves in their chartreuse skin are spiky and fresh. Further up the slopes, some of the blue oaks are sprouting. On the blue oaks I see, the sprouts are not at the bases of the trunks but higher on the tree, on the branches. These differences are characteristic of the two oak species: live oaks tend to regrow from their bases, blue oaks from their crowns. The oaks that are able to survive fires will benefit from them, especially if the fires were low intensity. Fire removes accumulated litter beneath the trees, which helps reduce the numbers of insect pests that feed on acorns, such as filbert worms and filbert weevils, two of the most common in California. Fire also helps remove some of the oaks' competitors for resources, such as annual grasses, though it can also open opportunities for other competitors\u2014thistles, filaree, and the like\u2014to gain a foothold and claim the open turf for themselves. The world underneath the blue oaks' canopy is much different now than it was before incursions of plowing, plants, and livestock. Fire can still be a healthy force, but that is no longer always the case, now that fires burn hotter and higher, and more often kill the crowns of trees, making it hard for trees to survive and resprout. The bark of mature blue oaks is thinner than other similar oaks, which increases the trees' susceptibility to fire as they age. It is more difficult for acorns to grow under competition from introduced plants and in our rapidly drying environment, with droughts becoming more and more common. All of this is to say that one year, three years, five years in, it is too early to tell how the oaks are doing at Cold Canyon. Trees challenge our impatient animal desire to know what is happening. Acorns take a very long time to make more acorns. Seedlings take a long time to become saplings, which take a long time to become mature trees. Time stopped for the branches frozen by the heat of the fire, but most of those trees are still alive and sprouting green at their bases. Trees live and die slowly, but it is difficult to remember this in the context of a disturbance like wildfire. The fire can feel like a catastrophe, and tallying the damage afterward feels so urgent. Trees move in their own time, and it is hard for me to comprehend when I am so quick, so mobile, so unrooted. Oak gall wasps Sometimes I go hunting for marvels, and sometimes they are sitting right in the middle of the trail, in shocking bright-pink glory. On a day when I arrive at Cold Canyon planning to focus on oak regrowth patterns, it is a strangely appropriate gift to find a crackly, dry blue oak leaf in front of me, covered in tiny galls. There two different kinds of gall on the leaf, both elaborately structured and very pink. I pick up the leaf and sit down on a rock beside the trail to wonder at the details of these tiny homes, nurseries to the microscopic babies within. The interaction that creates these intricate structures is an amazing tale of manipulation, and a relationship maybe even more strange than the complex interaction fostered by horntails, fungi, and the distressed trees in which they lay their eggs.* Once again, it is wasps that are responsible. This time, very much unlike the large, somewhat alarming horntails, however, gall wasps are no bigger than about a quarter of an inch. Some of the smallest gall wasps in the world are only one millimeter long, the size of a comma on this page. Female gall wasps lay their eggs in rapidly growing parts of trees, such as twigs or leaves like the one I am holding in my hand. Once the eggs hatch and the wasp larvae begin to eat the plant tissue, a chemical in their saliva, or perhaps the mechanical process of their feeding itself, stimulates the plant to redirect some of its own cells to produce a protective outer structure that surrounds the larvae. This is a defense response by the plant, sequestering the larvae so that they do not eat its other parts. But it has great benefits for the larvae as well. The gall is both shelter and, inside, more food for the ravenous larvae, which comfortably feed until they mature and chew their way out of the nursery as adults. It does not seem as though the wasps do any real harm to their host plants, though they do of course consume some resources that would otherwise be used to meet the plants' own needs. What is most amazing, and apparent even on the leaf I've just found, is that each species of gall wasp stimulates a very specific and unique structure of gall. One of the kinds of gall on the leaf has spines sticking out in all direction. This is the home of a batch of urchin gall wasp larvae. The other kind is almost furry, and is made by crystalline gall wasps. The gall shapes are even expressed in the wasps' scientific names. The urchin gall wasp is Cynips quercusechinus\u2014the species name means \\\"oak hedgehog,\\\" emphasizing its spikes. The crystalline gall wasp is Andricus crystallinus, inspired by the delicacy and translucence of its fine \\\"hairs.\\\" Two different species, with two characteristic gall shapes. And there are over a hundred different gall wasps known so far worldwide. How on earth do the chemicals in the saliva of the larvae dictate the shape of the gall that the plant cells will build? Most gall wasps are specific to a single type of tree, usually an oak. It turns out that blue oaks have the largest known number of different gall wasp species, at forty-one and counting. They also appear to have the greatest diversity of shape and color of galls. I am looking at the galls just a little after one year since the fire. I try to guess which of the blue oaks around me dropped this leaf. I wonder whether it was a tree that burned, and this was a new leaf since the fire, or one that survived the fire intact. Perhaps the wasps whose offspring were reared in these galls came to Cold Canyon after the fire, or perhaps they survived the fire as larvae themselves. The effects of fire on oak galls have not been extensively studied, but it appears that gall wasps need time after a fire to return. There must be oaks for them to return to, either newly grown oaks or ones that survived the fire. There must be sources near the burn for them to come from. And some wasp larvae surely survive the fire, sheltering in the oaks that did not burn and remained cool enough that their leaves were not all killed by the heat. Adult wasps are not likely to survive the fire themselves, but then, they generally live only about a week, so the wasp truly spends most of its life\u2014about a year\u2014as a larva. How funny it is that wasps have twice been my windows into the endlessly intricate webs of interactions that compose these ecosystems. First, the horntails, who rush to burned trees to lay their eggs and inject their symbiotic fungi. Now the gall wasps, who bend the oaks to their will but are dependent on their hosts for shelter and protection when the world around them burns. Without being present in these places\u2014burning and burned\u2014I would have remained ignorant of these beautiful mysteries. The wasps' relationships are just glimpses into vast silken webs of stories that humans have not yet even dreamed. Start traveling along just one exposed thread, and how many more nodes in the silk might we find? The ones we do see tantalize us with the many that are yet unknown. Gray pines: Waves of loss and return Gray pines, towering over the other trees at Cold Canyon, seem to me the embodiment of solidity and permanence after the fire. They have an underappreciated beauty, being arguably rather spindly and dull in comparison with some of the more majestic pine species. But I think their long, pale-green needles are pretty, especially in the bright summer sun. And while their needles are not densely packed enough to create a whole lot of shade, I find they create striking silhouettes against wintery overcast skies. In the early years after the canyon burned, I am anxious to count the survivors and the lost, to take inventory, even if anecdotal, of the fire's casualties. Studying the gray pines, I can see green needles on what look like two-thirds of them, so I assume they have survived fairly well. But on a winter hike with Jeffrey Clary and Sarah Oktay, the Reserve's current director, just over three years after the fire, I learn that the truth is likely more complicated. Gray pines, also known as foothill pines, are often found in the blue oak woodlands and savannas that are interspersed with chaparral habitats in the California foothills. Like blue oaks and chaparral shrubs, they are hardy and thrive in places with drier climates and less nutrient-rich soils. Unlike many other plants, they do not have root, trunk, or bark adaptations that allow them to survive fire. Gray pines are extremely pitchy, even for a pine. They are torches in a fire, thanks to the resin in their needles, cones, bark, and wood. Because they do not contain the burls or other root structures that allow many chaparral shrubs to regrow, a gray pine too damaged by fire will not grow back. Instead, gray pines must start all over again, growing from the seeds protected inside their enormous, spiny cones. I see cones, open and dry, with seeds long spilled, all over the canyon. Picking them up and turning them in my hands, I wonder whether their seeds are unfurling now, nestled underground where they landed after dispersing when the cone opened still high on its tree. The heat of fire stimulates germination by weakening or breaking the seed's coat and allowing it to begin development. Gray pine seeds are also able to germinate without fire, but there are other pine species, such as lodgepole and knobcone, that do require fire for germination, at least in some parts of their ranges. This all sounds reasonably straightforward: gray pines mostly die in fires and eventually return to the landscape as new sprouts. It is the timing of their dying that is a surprise. Jeffrey points out that even though many of the trees at Cold Canyon still have green needles in their crowns, they may already have been killed by the fire, either because part or all of them burned or because the heat of the fire was too great and killed their tissues. Set inexorably onto the path to their fate, they are continuing to function while slowly winding down. In their diminishment, they are now home to organisms that they would have easily fought off while healthy. The fire drew pine sawyer beetles\u2014a kind of longhorn beetle\u2014and their larvae are now tunneling through the pines' wood. Bark beetles, such as pine engravers, will also have come, finding nourishment in the wood of the dying trees. These wood-boring beetles and their relatives are usually healthy members of forest communities, consuming only trees that are already dead or dying and returning the trees' nutrients to the ecosystem. But in the increasingly stressed habitats of the western US, some species, such as the pine engravers, have become decidedly unhealthy. In the face of plentiful dead wood during years of drought and fire and excessive logging, the beetle populations explode. Voracious new generations feast on living trees too, making for even more fuel in wildfires and escalating the vicious cycle. While we are most familiar with large expanses of dead pines and firs in the Sierra, gray pines are not immune to these amplifying effects of climate and beetle. As I walk beneath a pine with silvery-green needles left on only a few of its higher branches, I see the fine line between health and disease, between beetles as beneficial recyclers and beetles as forest destroyers. I am learning that dying is a completely different phenomenon for a tree than for all of us more ephemeral creatures. It is not a single event in time but a very long process. How smooth is the continuum of living and dying and how hard to say where some organisms are on that spectrum.* There are no quick answers in assessing the results of upheavals like wildfire, at least for trees, as I watch the aftermath play out slowly over the years. Gray pines are common in chaparral habitats, but they require plenty of time to recolonize between wildfires, after the fires have mostly killed off the existing trees. On the one hand, if wildfires occur too frequently in chaparral, there will not be enough time for new gray pines to grow from seed. As the window for recolonizing closes, gray pines will slowly disappear from these habitats. On the other hand, if fire is completely absent from the chaparral ecosystem, gray pines will take over\u2014enough gray pines will eventually colonize that the blue oaks cannot compete for space and sunlight. Fire\u2014at least a healthy fire regime\u2014keeps the balance in these blue oak and gray pine foothill habitats. After my hike with Jeffrey and Sarah, before I head home to the valley flatlands, I stand next to a gray pine and put my hand on its bark. Time slows to a crawl as I deliberately calm my quick animal breaths. Time stops while I try to think in tree time. Here time is measured in roots creeping, tunneling, and connecting with fungal mycelia and other roots. Time passes in rings of wood expanding, thickening, drying, and cracking into bark. And time is the slow, slow pull of senescence, embarked on long before the tree's final end. Senescence is set to the tiny music of the jaws of beetle larvae scraping and gnawing their intricate way through the tree, just below the surface of the bark. This is a kind of sorrow, deep and abiding. There is nothing that will stop the onward pull of time and the tightening of death's grip. The last green needles will brown. The wood will fully dry. The tree will continue to stand for a long time, entirely dead, waiting for the gust of wind or shift of soil that will send it crashing to the ground. But I am also, just as deeply, moved by the cycles in which the pines and I are enveloped. As though my awareness of the process allows me into the rolling pattern. Nothing is what it once was, nothing in the view ahead of me can stay this way forever, and we are all senescing even as we are full of life. The gall wasps whose tiny homes I marvel at on blue oak leaves live only a week as adults. A blink of an eye. What would they make of my ponderously long and slow life, beyond the limits of comprehension? Just as I struggle to understand how a living tree can also already be dead. * For more on how trees blend of life and death, see Peter Del Tredici, \\\"The Roots of Rejuvenation,\\\" in this issue (page 30). robin lee carlson is a natural science writer and illustrator based in Davis, California. Visit her website at robinleecarlson.com"},{"type":"arnoldia","title":"Morus rubra","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25876","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e3d25ebb6a.png","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"4","authors":"Jaikumar, Devika","start_page":"14","end_page":"15","article_content":"In the summer heat, mornings are blanketed by a mist that leaves the arboretum quiet and still. I softly make my way up Bussey Hill, my shoes soaked through by the morning dew that lies thick on the grass. A fawn suns itself on a patch of dirt while a family of turkeys take their morning stroll, all undeterred by my presence. It isn't these creatures I have come to see, however, but 787-85*A, a striking mulberry tree that every morning for the past two weeks had been picked clean of fruit by squirrels. Found all over the world, mulberries are known for their sweet fruits that resemble blackberries, and for their role as a key food source for silkworms. The tree I visited nearly every day for a month is a specimen of Morus rubra, a species native to North America. At the Arboretum it is surrounded by Morus alba trees, a species introduced from Asia. Despite their differences\u2014glossiness of leaves, presence of hair-like structures on the underside of leaves, colors of fruit\u2014these species are prone to hybridization, making positive identification of trees especially complex. My goal in the summer of 2021 was to acquire a cutting of the plant, with leaves and fruit, to press for an herbarium specimen. As I approached the tree that morning as I had so many times before, the squirrels leaped off, revealing that they had picked the branches clean of ripe fruit. I could not fault them for their behavior; the idea of explaining to the council of squirrels how I needed a handful of fruits for scientific study was enough to keep my spirits up until the day I was successful in my task. The specimens that I was finally able to collect were sent to Morris Arboretum, where researchers are engaged in long-term study of mulberry hybridity. Herbarium specimens have been crucial to research for centuries, and one of their primary uses is to identify species. While we believe our multi-stemmed Bussey Hill tree is correctly identified as Morus rubra, perhaps it is indeed a hybrid, the identity of which has taken years of growth to reveal itself. Herbarium specimens also fill gaps in our knowledge by creating a physical timeline for plants throughout their lifespans. When I collect a specimen to later mount onto paper and file into the Arnold Arboretum's Herbarium of Cultivated Plants, it's with the knowledge that this specimen is intended to last hundreds of years, to be used in ways that I can't even imagine. As for the squirrels, their interest in the mulberries is far less mysterious."},{"type":"arnoldia","title":"The Hidden Ice of Plants","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25805","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060b76a.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"3","authors":"Villouta, Camilo","start_page":"11","end_page":"12","article_content":"When studying how plants in winter deal with freezing temperatures, the observation of ice formation in their tissues is especially elusive. I addressed this topic in my dissertation, which challenged me to find new methods to answer them. At the time, I was a PhD student at the fruit crops lab at the University of Wisconsin-Madison, under the guidance of Dr. Amaya Atucha. Our goal was to understand the development of cold hardiness in the terminal buds of cranberry (Vaccinium macrocarpon). Most importantly, we wanted to know what strategy they used to withstand and survive ice formation. In woody perennial flowering species, buds are known to survive exposure to freezing temperatures by either of two freezing survival strategies. Both follow a similar principle: avoid the formation of ice crystals with their potential of causing lethal damage in the flower primordia, tissues that are the precursor of the mature flower. These strategies have extravagant names, which nonetheless describe their main characteristics very well. On one hand, we have \\\"deep supercooling,\\\" where buds avoid the propagation of ice into the flower primordia, keeping them ice-free even when they are at subfreezing temperatures. The other strategy is named \\\"extraorgan freezing,\\\" where the flower primordia dehydrate, pushing water away from themselves towards the ice-tolerant bud scales. In the case of cranberry terminal buds, we needed to determine which strategy was at work, both to interpret damage patterns more accurately and to know which techniques to use for assessing cold-hardiness levels at different times across the season. While designing the study to answer this question, we reviewed a range of current methods for imaging plant tissues, assessing them for their ability to capture freezing events inside an intact bud. Thermal video recording offers only a surface view of freezing progression, as do electric thermometers, or thermocouples, which only register temperature changes at a specific location. MicroCT (computed tomography) scans offer great resolution of internal tissues, but cannot track freezing, as the method does not detect a contrast difference between liquid water and ice. Magnetic resonance imaging (MRI), however, can detect this difference. Searching the UW campus for appropriately-sized MRI machines, I found one in use at the Small Animal Imaging Center. While a regular MRI has an entrance designed to fit a person, this apparatus was made for animals such as rats, with an entrance diameter of about 3 inches. Once we found the proper MRI machine, we needed to find a way to control the rate of decreasing temperature at which the buds would have been exposed\u2014a rate of vital importance. Studies have reported that a common rate of temperature change in nature is 1 \u00b0C\/hr (degrees Celsius per hour). Thus in laboratory conditions, we usually work with rates ranging from 1 to 4 \u00b0C\/hr, otherwise you risk creating artificial effects in non-realistic freezing conditions. To detect the progress of freezing in our plants, I needed a device to control temperature with great precision, which I set out to construct. The goal was to acquire MRI images corresponding to slices of our samples at room temperature, and then slowly decrease the temperature, capturing other images at just below freezing, and then two more times at colder temperatures, always of the same slices. In collaboration with the Morgridge Institute's Advanced Fabrication Laboratory,\u2014 a.k.a. the \\\"Fab Lab\\\"\u2014at UW Madison, we started to develop a prototype. It needed to follow several guidelines: it could not have any of its metallic implements close to the MRI apparatus, while at the same time it had to withstand freezing temperatures, exhibit chemical resistance, and not interact with the imaging process. The prototype needed to have three compartments: one for the flow of chilled glycol, a second for the addition of a chemical that helps with image contrast, and a third to surround the samples with all these compounds. Once the design was defined, we started testing the circulating system. We located the circulating bath and pump for the chilled glycol with their metallic parts outside the room, connecting the glycol via tubing with the prototype inside the MRI. With this circulation system exposed to air, frost formation was also a big concern, as this could damage the expensive MRI equipment. We ran trials in our lab just to see where frost was forming and find corrective measures. By this time, a year had passed, and finally, we were able to obtain our first images. After doing the official runs in late fall of 2019, I measured the signal intensity at different regions of the buds and compared them across the different imaged temperatures. Higher signal intensity translated into a higher presence of liquid water, and the opposite meant relocation or freezing of water. My analysis showed a gradual decrease in the amount of liquid water in the internal tissues of the bud. This was an important piece of evidence for our study. With the MRI, we observed a range of freezing events occurring at temperatures below -20 \u00b0C, while in visual damage evaluations for that same date, we found that damage did not start until reaching temperatures of -24 \u00b0C and lower. In the end, our study concluded that terminal buds of cranberry survive exposure to freezing temperatures by undergoing a process of freeze dehydration, a variant of the extraorgan freezing survival strategy. From here, we can direct our efforts on developing cold hardiness models with a greater understanding of the damage patterns, knowing what mechanisms need to be developed during the fall during the seasonal acclimation to winter. This knowledge will help farmers and horticulturists decide how to respond when fruit crops are exposed to the threat of unseasonably cold temperatures\u2014conditions likely to occur with greater frequency as climate change advances."},{"type":"arnoldia","title":"Bud Morphology","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25806","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060bb6d.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"2","authors":"Moeglein, Morgan","start_page":"8","end_page":"10","article_content":"Every fall across the temperate latitudes, the leaves of woody plants complete development, reach their final form, and begin to senesce. Leaf senescence marks the end of a growth cycle that may seem to begin at bud break in the spring, but for many leaves actually began much earlier, during the previous growing season, hidden from view within a resting bud. Leaves that initiate and mature within a single growing season are known as neoformed leaves. Preformed leaves, by contrast, initiate during one summer, undergo one or more periods of winter dormancy, and complete their life cycle in the following summer. Botanists have long noted the presence of incipient leaves within dormant resting buds. Many unanswered questions about leaf preformation remain, however. When we look at a tree at the end of the summer, how many of those leaves were preformed? Do different species employ different strategies when they invest in preformed leaves for future seasons? I have spent the last year trying to characterize how plants invest resources in their overwintering buds and the leaves within. This work led me to the Arnold Arboretum, where I set out to investigate how preformation contributes to leaf development across woody taxa. The first step in this work was to look very closely at dormant buds in the field. Surveying buds across species in the depths of winter, without the obstruction of leaves, is the best time to appreciate how much bud morphology varies. Some overwintering buds are obvious, such as the long, shiny beech buds or the furry, gray magnolias. Then there are the minuscule and hard to find: the buttonbush bud barely raised from the surface of the twig, or the winterberry with its few, tiny, camouflaged bud scales. Walking through the collections sampling dormant buds for future microscopy, I easily snapped the bigger buds from their branches with my fingers, while smaller specimens had to be shaved from twigs with a razor blade back in the lab. Even more remarkable than the external bud morphology observed in the collections is what you find inside after dissections under the microscope: the large, showy buds usually contain mature-looking leaves in miniature. The morphological intricacies of an oak or sweetgum leaf are already apparent, intact in these preformed leaves, months before they expand to their mature size. Even in the smallest buds, many leaves are already initiated, their shape recognizable in its relation to the mature form. A tiny, flattened catalpa bud may contain ten leaf primordia, similar to the number of mature leaves that will be present at the end of the growing season. It's astonishing to realize, while looking at buds in the field, that the form and number of mature leaves could be determined so far ahead of time, within the tiny space of the smallest buds. By comparing preformed leaves dissected from within buds to mature, fully expanded leaves, I hope to deduce how much of the multi-seasonal cycle of leaf development occurs within buds relative to the end-of-season total. Looking at preformed leaf and bud measurements relative to mature leaf measurements allows us to measure relative investment in different leaf components, and begins to reveal patterns in preformed investment. These strategies can tell us something about how plants prepare for the future. Some species may preform more leaf tissue that is more fully developed in preparation for expected circumstances in the coming growing season, while others may wait and neoform the leaves they need in real time as a response to present growth conditions. There are also different ways to invest. One species might invest in many small leaves, while another might invest in fewer leaves that are larger and more completely developed. With so many different ways to invest in preformation, the range between preformation and neoformation begins to seem more like a spectrum than a dichotomy. Based on observations so far, I suspect the species surveyed will vary widely in the timing and extent of their investment in future leaves. Surveying the contents of overwintering buds can help us learn how plants invest in leaves from season to season\u2014but it may also better our understanding of the evolution of leaf form more broadly. Leaf shape varies massively across plants, but the variation within species or within individuals can be just as impressive. Leaf shape along a single growing tip can vary over the course of a season and this variation may correspond to preformed and neoformed trajectories arising from different developmental programs. Leaf-shape plasticity could provide standing variation for evolution to act upon, or it might open up different photosynthetic options as the season progresses. If the shape of preformed leaves is largely predetermined within the dormant bud, and if this shape is different from the shape of neoformed leaves later in the season, the confined nature of development within a bud may have some influence on leaf shape. The same may be true when comparing across species with different preformation strategies; a species that preforms more may be more likely to have certain leaf shapes than a species that produces more leaves through neoformation. There are many possibilities, but looking within dormant buds at the earliest stages of leaf development across species and environments may illuminate our understanding of the variation we see across mature leaves. IN MEMORIAM Karen Madsen 1942-2022 Karen Madsen edited Arnoldia from 1993 to 2007, leading the magazine through dramatic changes in publishing and printing. Perhaps no one at the Arnold worked with Karen more closely than Peter Del Tredici, who offers the following remembrance: I first met Karen in 1988 when she was a student in the Radcliffe Seminars program, working on her thesis on Woodlands, the famous garden of William Hamilton, who introduced the Ginkgo tree into North America in 1784. When I was appointed editor of Arnoldia in 1989, I asked Karen to contribute a short version of her thesis. We got along well during the editing process, which is not always a given, and I hired her part time to help me with copyediting. After I became Director of Living Collections in 1992, Karen was appointed to replace me as editor. Not only was Karen a good friend, a great writer, and a skilled editor, but she helped make landscape design history one of Arnoldia's specialty topics. BOOK BRIEF Older Than the World \\\"Trees,\\\" writes historian Jared Farmer, \\\"are plants that people call trees\u2014a term of dignity, not botany.\\\" And it is the largest and, especially, the eldest that have commanded the greatest dignity, from the the Buddha's Bodhi Tree to Methusaleh, the bristlecone pine in California's Inyo National Forest thought to be the oldest non-clonal tree in existence. Believers held that the Oak of Mamre in Jerusalem had been growing since the dawn of creation\u2014a tree older than the world itself. By the time this tree fi nally fell in the yard of a West Bank monastery, far older trees had been identifi ed by scientifi c means. And yet fascination for trees like the giant sequoia accompanied a rush to cut them, nearly leading to their disappearance. Farmer's book explores the mythmaking, veneration, and exploitation to which the oldest and largest trees have been subjected through time, and poses an urgent question: in the wake of the industrial era, under the cloud of climate change pushing them to the brink, do these elder fl ora have a future? We need them to survive, Farmer argues, if we are to make sense of changes we have set in motion in deep time."},{"type":"arnoldia","title":"Converging Paths","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25867","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e3d24e816d.png","year":"2022","series":null,"season":null,"volume":"79","issue_number":"4","article_sequence":"1","authors":"Battles, Matthew","start_page":"1","end_page":"2","article_content":"Two roads diverged in a yellow wood: Frost's words come to mind unbidden as I walk through the Arnold amid November's yellowing days, following forking paths that might take me to the Hunnewell Building and the workday, to Weld Hill for a lecture, or into the conifers for a quiet afternoon copyediting in the company of wild turkeys. It's worth remembering that intersections are also convergences, comings-together for travelers from different points of origin. The arboretum itself is such an intersection, where two grand ways of working with plants, the scientific and the horticultural, converge. How many other ways of knowing, feeling, and storying trees also meet at this crossroads? Some convergences come after long travel. Turning off Meadow Road, I approach the golden glow of the katsuras\u2014though even before I arrive, they've already announced themselves by their characteristic caramel fragrance. One of these specimens of Cercidiphyllum japonicum, 882*A, is among the oldest trees in the arboretum, accessioned in 1878. But katsura's path is far older: fossil evidence shows them thriving in forests of the northern hemisphere by the end of the Cretaceous, some seventy million years ago. We say that katsura smells like cotton candy\u2014 the German name for the tree is Kuchenbaum, or \\\"cake tree\\\"\u2014but isn't it properly the other way round? Cakes and cotton candy, those latecomers, smell like katsura, whose fragrance of toasted sugar, tens of millions of years in the making, likely wafted long before there were noses attached to humans with a propensity for naming, propagating, and choosing tree-convergent ways. So these converging paths through trees are generous, gregarious, inspiring. Richard Powers begins his 2018 novel The Overstory with a powerful invocation of trees as generous storytellers: A woman sits on the ground, leaning against a pine. Its bark presses hard against her back, as hard as life. Its needles scent the air and a force hums in the heart of the wood. Her ears tune down to the lowest frequencies. The tree is saying things, in words before words. It says: Sun and water are questions endlessly worth answering. It says: A good answer must be reinvented many times, from scratch. It says: Every piece of earth needs a new way to grip it. There are more ways to branch than any cedar pencil will ever find. A thing can travel everywhere, just by holding still. The woman does exactly that. Signals rain down around her like seeds. With this year's Spring issue, Arnoldia debuted a new design and editorial format; in the very same moment, I joined as editor. I can take no credit for this new vision for the magazine, which was brought forward by Michael Dosmann, and my predecessor, Jonathan Damery, in collaboration with members of Arnoldia's editorial committee and our consulting design studio, Point Five. I've been thrilled by all the new ways they have furnished me to know, story, and connect with trees\u2014not only through a profusion of new literary forms, including poetry and creative nonfiction, but drawing from the visual arts as well. We've welcomed verse from emerging and prizewinning poets, and book excerpts from provocative scholarly works like Rosetta Elkin's Plant Life (Fall), to Cold Canyon Fire Journals by Robin Lee Carlson, whose distinctive, lyrical art and storytelling about wildfire and place in California enriches this issue (page 16). Our editorial artist, Matt Huynh, provides each issue's Notes from the Field section with a fresh set of expressive illustrations, while our endpapers by Shyama Golden frame the magazine with sumptuous assemblages drawn from the shifting seasons. At the same time, this renewed Arnoldia also brings forth the scholarly and scientific research that have long been the magazine's mainstay, such as Peter Del Tredici's article in this issue on the fascinating phenomenon of endocormic roots (page 30). And most crucially, we're making space for authors whose voices historically have not been heard along the paths of science and the garden. Native voices, the voices of folks of color, the unhoused\u2014Arnoldia seeks to come into restorative relation with such authors, and to bring the knowledge they carry to our readers. Arnoldia's rebirth comes at a time of both richness and urgency, with an efflorescence of innovative writing about the natural world arriving from new directions, prompted by climate change, calls for environmental justice, and the enormity of the biodiversity crisis. And so now is a crucial time to bring forth this expanded editorial mission and toolkit for telling tree-entangled stories. Meanwhile, the Arnold's paths proliferate. The horticulture team has been laying new trails throughout the landscape, giving visitors more immediate access to our collections. And in the years to come, the Roslindale Gateway Path Project will connect Forest Hills to Roslindale Center, making even more of this living landscape into an equitable contact zone. As these paths intersect, new stories surely will emerge. Our former tagline was The Magazine of the Arnold Arboretum; now, it's The Nature of Trees. An intersection for more than science and horticulture, this nature is a gregarious convergence of lyric, memoir, storytelling, and scholarship. Who speaks to the nature of trees? Everyone. May Arnoldia continue to be as generous as the katsura, as message-bringing as Powers' fragrant pine. May it be a crossroads, a meeting place. And may it always be thronged with trees."},{"type":"arnoldia","title":"2022-79-3","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25793","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15e816e.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","arnoldia_cover":true},{"type":"arnoldia","title":"Hardwood Cuttings","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25808","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d0608528.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"14","authors":"Halloran, Sean","start_page":"64","end_page":"64","article_content":"Plant propagation is both traditional art and applied science. Made from mature, dormant stems, hardwood cuttings are just one clonal propagation technique we use at the Arnold Arboretum to coax life from some of our most valuable accessions. In New England, we can root hemlock and many other genera via this technique. We collect hardwood material in late fall or early winter after plants enter dormancy and deciduous plants have lost their leaves, selecting younger, healthy stock for optimal results. Hormone is applied to the freshly prepared cuttings, which are usually \\\"stuck\\\" the same day they are collected, in benches filled with quick-draining growing media, tented with plastic to ensure high humidity, and heated from below to encourage rooting. Hormones and wounding treatments are taxon-specific, based upon the scientific literature as well as our own propagation records. Cuttings that survive and root will be potted up in the spring or early summer, eventually to serve as prized members of the next generation of our collections."},{"type":"arnoldia","title":"Among white pines, at the foot of them","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25809","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060856c.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"13","authors":"Lorsung, \u00c9ireann","start_page":"61","end_page":"63","article_content":"The largest city in the place I call Maine is called Portland, and it lies along the Atlantic coast in the northeast of the country called the United States. For as long as the city has existed its industries have been tied to the sea: fishing, lobstering, clamming, oystering, seaweed-picking\u2014though that is changing: wealth clusters along the seafront and the city's industry turns toward tourism. At low tide the horizon goes even further out than you'd think possible, and between the expensive houses and the drawn-back water, the seaweed pickers go, bent over, along the little strip of land that belongs to everyone. The coastline just north of the city becomes rocky, and small granite islands house wealthy summer residents and year-round island people who know the ferry schedule by heart. A map eye and a surveyor's eye, a deed eye; an insurance company's eye, a property investor's eye, a painter's eye; an art collector's eye beholding the paintings of the \\\"empty\\\" landscape that hang in the well-appointed houses in the best neighborhoods, in the white cool rooms of the city's art museum. We perceive the beauty of a map or painting in the manageable space between the object and ourselves, the gaze a place that holds the object still. In the map and the landscape painting\u2014 objects I feel such a fondness for\u2014the place that holds me is reduced to a place that I can hold. It becomes property and I am its arranger. The world goes on around the museum and the city records office, in infinite overlapping circles. The room and the building, the city and the state, are permeated by economies and public-health statuses and modes of transit and pollution, by taste and custom and the determining structures of US racial and class hierarchies. We work temporary jobs in Portland and pay two-thirds of our income to the landlord. It is a pandemic, so we don't go anywhere. We couldn't afford to go out in this city in any case. To occupy my time and thoughts, I take walks and ride my bicycle: watching the light change is free, noting down the seasonal appearance of weeds in the cracks is free. I am moving through a landscape that is unfamiliar to me, and I cannot see it from outside\u2014outside the drafty, unheated apartment, outside the exorbitant rent, outside the virus transmitted by breath. Nor from outside the sea air, the smell of the white pines, the sting of first snow on a stiff breeze. Only from inside, where I am: with you. I find my way, in copper October, along a causeway to an island that houses the state school for Deaf and Blind students, a cemetery where the dogs and horses of a former governor have long since turned into soil themselves, and a clearing in which visitors build structures out of twigs, leaves, shells, seaweed, rocks, pinecones, and a winking belief in fairies. (The children who visit to see the fairy village do not wink, and their houses are furnished with bright orange berries, mushrooms, moss carpets, colorful and smooth stones, the best shells.) The name I know for this island is Mackworth. Among the trees, the name I know falls away, insubstantial as a piece of Scotch tape. Its granite edges, its carpet of rust-orange pine needles, its non-local roses trimmed back by state park service employees, its sumac, its dulse. I have names for these, and yet the names cannot hold them. They exceed my ability to call them. In every season, the island's white pines establish the architecture of my passage there. I walk the circumference in autumn's fullness, then in the silence of winter as the wind stirs the ocean into gray slush, and in spring as a bright yellow haze passes through the deciduous trees of the understory. And in the summer, in the rich quick bright unbearable green of living things that know how soon they will be dying. The tall gray bodies of the pines set a rhythm for seeing. Their needles form paths and fields among the trees. The sunlight falling through them, I receive. The sky they orchestrate among their openings. They are an uncapturable color, a green that is not singular green but green-prismatic with all earth colors: chlorophyll, umber, ochre, iron oxide. Light and vision refracting in air. Among the white pines, at the foot of them, I find a distinct understanding of my proper proportion. The placedness that makes me among and with and makes my gaze through and in rather than at. From the edge of the city I see them, objects on a horizon: but among them, my sight is transformed and I am a subject among subjects. I would like to hold them in this withness and throughness. \\\"Landscape\\\" is an effect of an exterior position. I am wrestling with my love of landscape painting and my position as an outsider to that tradition, by virtue of gender, training, and class\u2014and I am wrestling with the fact of my being-settler in this place, with the history of ownership- as-land-relation I belong to. From a distance, the painter or surveyor perceives an interplay of light, shadow, form, movement. These are arranged as painting or as map, or, if in the mind rather than in material, as property. But this island is a freely offered immersion in color, perspective, time. Surrounded by white pines, it is impossible to take distance. I cannot stand back and see myself apart from them. They remind me we have always been involved in one another, whether I knew it or not. I was formed in a scientific and religious environment where human beings and other beings are unequal. But despite how my imagination or understanding was formed, the white pines' way of being requires, structurally, that I am among them, beneath them, between them, in their midst. It changes the kind of relationship I can have, and not just to the white pines. The withness they do to me extends limitlessly. They with me, and they with their understory. They with the greyness of the ground, scattered with their rust-orange needles, after long weeks without rain. They with the chipmunks who make holes at their feet. They with and are withed by the non-local roses and the local ferns, the lichen, the sound of the tide arriving, the bladderwrack, the smell of the mud on a hot afternoon coming through the closer sweetness of the needles that carpet the dry ground. They with the conditions of the air\u2014humidity, pollution\u2014and of the water, weather, the salt marshes, the bogs beyond, the fields beyond that. The way the white pines are makes me with, a with that exceeds them and far exceeds me\u2014and exceeds the pronoun \\\"me\\\". Here, among the trees, I find myself one among many, even unseen many, moving in a concert it is impossible to know in full. The island, the ocean, the iron streaks in granite, the yellow seaweed and its scent in the September air, the red hips of the roses spattering the afternoon: I don't want a painting I can own as soon as I see, that lets me say I know what you are. I want something that can do to me again what has been done here\u2014that amongs me. As color does to me, as scent does: throughs me. Withs me. Bewiths me. This human head and your lowest branches. The light held there, having fallen through you from the nearest brightest star. The air I expel from my lungs and take into my lungs. The words-failness of that light, air, open, up, around space. An experience closer to what I name preposition than noun: more than thing it is movement, direction, location, a totality of space and passage. The pines make this sense as they move through time and space. By the time I arrive where they are, they have ordained an entire way of being, and I enter it. This is true of almost all places I go: the city and its apartments, the museum and its rooms. They have been made in advance and they will outlast me. Among the white pines on this rocky island I am altered to fit the time and space of the pines. What is this alteration? It changes the I that I am, and the eyes that I have. The duringness and withinity of the pines are a grammar that refits me to an accompanied and accompanying multiple subjecthood. No longer the center of the sentence-shaped universe or the distant viewer of a landscape made to hold, I am here: with the pines, with the world they are."},{"type":"arnoldia","title":"Plants, Identity, and War in Ukraine","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25810","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d060896f.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"12","authors":"Keegan, Brendan","start_page":"58","end_page":"60","article_content":"From 2010-2012, I served as a U.S. Peace Corps Volunteer in a small Ukrainian town. In 2017, my wife (who also served in Ukraine) and I returned for a visit. We met with Natalia, a dear friend, and former colleague, at the school were I once worked. Together, we walked the old paths connecting nearby villages through the surrounding forest. It was a crisp, September day. The previous year's leaves, from towering old lindens, crunched beneath our feet. As we walked, I remembered the forest in seasons past. In March and April, millions of blooming white wood anemone and snow drops carpet the understory as far as the eye can see. A neighboring baba (granny) once asked if I would pick her a bouquet of those fragile, ephemeral flowers. As a child, she had plucked from those same woods. Giving them as gifts is a tradition common in the region for good luck and health. Last spring, I observed Arboretum visitors marveling at the same delicate snowdrops popping throughout our own collections. I thought of that friendly baba. As news images show the utter destruction of Ukrainian cities, towns, and villages, I often think of her, long since deceased, and her love for those simple flowers. Many readers will surely appreciate the Ukrainian custom of sharing snowdrops, of marking the seasons with walks through old forests. Indeed, Ukrainian identity as a whole is strongly associated with the natural world, and with plants in particular. The kalyna, better known in the U.S. as a guelder rose or snowball viburnum, is the most prominent example. Ukrainians believe planting one outside a home brings health and good fortune. Its red berries also serve as a metaphor for Ukrainian nationhood and independence. The plant prominently features in Ukrainian music, literature, and art. Its red berries adorn the bright embroidery of the vyshyvanka (Ukraine's national costume) as well as the modern-day insignia of Ukraine's armed forces. Plants play important roles in many other aspects of the culture. Wheat, the nation's economic lifeblood, is famously referenced in Ukraine's flag, golden under a bright blue summer sky. European aspen, found at the Arboretum on Peters Hill, features widely in Ukrainian poetry and literature, especially in the work of Ukraine's national poet, Taras Shevchenko. In Kyiv, Ukraine's capital, horse chestnuts adorn the various squares; at one point in its history, horse-chestnut leaves even graced this green city's official seal. Botanically inclined readers might even be envious of Ukraine's calendar, in which the names of months reference coinciding natural phenomena. For example, Berezen' (March) means \\\"birch,\\\" and indicates the time to tap the birches for their sweet sap. Kviten' (April) is \\\"flower,\\\" when the earliest blooms appear. Lypen' (July) is \\\"linden,\\\" marking the long summer evenings when this tree's sweet aroma bathes Ukrainian towns. My personal favorites mark the fall. Zhovten' (October) means \\\"yellowing,\\\" referring of course to changing leaves, while Lystopad (November) translates literally as \\\"leaf fall.\\\" Examples such as these abound in Ukrainian culture. In addition to sharing a general love for plants, the Arnold Arboretum also shares connections with Ukrainian botanical gardens and their staff. During the first year of the pandemic, our virtual lectures and educational presentations provided an opportunity for a truly global audience to attend and participate. Staff from Ukraine's M. M. Hyrshko National Botanic Garden in Kyiv were among them. After communicating first over social media, then directly, they sought advice from Arboretum staff on plant curation and exchanges. Of course, everything changed for our Ukrainian horticultural counterparts when Russia invaded on February 24, 2022. As the Russians advanced towards Kyiv, the M. M. Hryshko National Botanic Garden closed to all visitors. While many Kyivites fled the city in the face of bombardment, Arnold Arboretum staff kept in touch with our garden connections who remained. Throughout the months of fighting, horticulturist Olga Pokhlychenko updated us on the situation in city and garden alike. As many a gardener can surely relate, she feared for her plants as well as her colleagues, friends, and family. Of course, she was not alone; a select group of staff were allowed into the garden to keep their treasured greenhouses functioning during the bitter February cold. When the garden closed, Olga joined multitudes of other citizens volunteering in defense of their city and for each other. She spent much of her time gathering and dispersing medical supplies, and later served as a volunteer coordinator. At one point, she darkly joked that she had so many medical supplies in her house that she had retired as a horticulturist and become a pharmacist instead. When the Russians retreated from Kyiv in late March, Olga and others went to assist civilians living in the formerly occupied suburban towns and villages. Several towns, including Bucha and Irpin, were off limits as a result of mines and booby traps left behind. However, in the numerous surrounding villages, they delivered aid to the disproportionate number of elderly who remained during the fighting. Some volunteer aid deliveries included seeds for vegetable gardening, a popular pastime and food source for both urban and rural Ukrainians even before the war. Even after the Russian withdrawal, the danger of attack remained so high within Kyiv that the M. M. Hyrshko National Botanic Garden stayed closed for several months. As winter turned to spring, many loyal visitors mourned the inability to see their beloved plants. They inquired about the status of their favorite magnolias, cornelian cherries, and forsythia. Unfortunately, even the garden's renowned lilac festival was canceled for security concerns. Finally, after being closed for over ninety days, the M. M. Hyrshko National Botanic Garden reopened for staff and visitors on May 28. The opening day aligned with the weekend-long celebration of Kyiv's city day, marking its 1,540th anniversary. Olga, and those of her colleagues who remained in Kyiv, finally returned to their plants. At the Arnold Arboretum, our own Lilac Sunday returned in full after two years of muted festivities. In tribute to our Ukrainian colleagues, we highlighted a rare lilac cultivar in our collection, Syringa vulgaris 'Ukraina' (pronounced oo-kray-EE-na), named in honor of Ukraine. Although still a small plant, our Syringa vulgaris \\\"Ukraina\\\" bloomed beautifully this year. The cultivar was discovered in 1974 by Ukrainian horticulturist Valentina Zhogoleva. She worked at what is today Ukraine's National Botanic Garden, where Olga and her colleagues continue her care for the lilac collection. A woman in a field largely dominated by men, Valentina is also credited as a collaborator on many lilac cultivars. All of her credited lilacs are named after Ukrainian heroes, cities, and literary icons. The Arboretum plans to introduce several of these now rare Ukrainian cultivars into our collections over the coming years. We also hope to share acquired specimens with Ukrainian institutions, such as Harvard's Ukrainian Research Institute, which may not already have them. Our hope is to provide plants for memorial plantings, remembering Ukrainians who have died in the ongoing war, while also preserving unique cultivars of Ukraine's botanical heritage. In early spring, I connected again with my friend Natalia. She provided updates on the town, my colleagues, and both the strain and unusual rhythm of war. As a teacher, she described the difficulties of conducting lessons with periodic intermissions in the bomb cellar. Like many of us, she found solace working outside. \\\"(L)ife finds its balance even during war,\\\" she wrote in her message. \\\"We also have spring here\u2014it is still cold, but tulips have already blossomed, we have a lot of fruit trees, so I have enough work to do.\u2026 And a rose flower garden of 52 rose bushes. My hands are all in thorns\u2014but it's not so bad. The beauty of flowers is worth it.\\\" I am sure that many readers would sympathize with Natalia's words. Perhaps peace is like Natalia's roses, painfully, and painstakingly, cultivated. Until it is achieved, I hope the shared love of plants, such as the beautiful snowdrops in spring, the flowers of our Ukrainian lilac in May, and the red berries of the kalyna each fall, remain symbols of personal and institutional connection with the Ukrainian people."},{"type":"arnoldia","title":"Public Gardens and the Livable City","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25811","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070a327.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"11","authors":"Rakow, Donald A.; Gough, Megan Z.; Lee, Sharon A.","start_page":"54","end_page":"57","article_content":"Over the centuries, public gardens like the Arnold Arboretum have welcomed visitors to their grounds and enhanced those visits with opportunities to learn about the garden's plant collections and landscapes and to reflect and relax. In recent decades, many gardens have expanded the garden experience through art exhibits and performances of musical and theatrical groups on the grounds. However, most visitors to public gardens are those with the means to travel to the garden and the resources to pay for admission, when a gate fee is charged. For residents of underserved neighborhoods, even public gardens that offer free admission may be largely unknown due to a lack of public transportation, the absence of bilingual signs, or a perception of the institution as unwelcoming and catering only to higher- income or white members of the community. To thrive in the twenty-first century, public gardens must respond to their elitist reputations and invest in programs and policies that are inclusive, caring, and relevant to a wider audience. No matter their size or budget, it is time for public gardens to assume their responsibilities as community institutions and to focus on outreach to all community residents. As plant-based scientific and educational institutions, public gardens have the expertise and the experience that can help address challenges such as unsafe neighborhoods, poor quality science education, limited access to fresh and healthy foods, a lack of job training and job opportunities, and degraded environments. In support of the role that gardens can play in addressing these challenges, studies have shown that as streetscapes become greener, they attract more activity and engender more pride in public spaces. When young children are provided with hands-on science and nature instruction, they are more likely to succeed in school and see science as a possible career alternative. When previously incarcerated individuals are given training and offered jobs in the green industries, they are far less likely to return to prison. And when under-resourced residents are given the tools to build and grow community gardens, their diets and health also improve. The most successful initiatives arise through partnerships. As we demonstrate in our book, Public Gardens and Livable Cities (Cornell 2020, foreword by Scott Medbury), many initiatives to improve quality of life in our cities result from partnerships between gardens and other community organizations, neighborhood groups, municipal agencies, and private entities. There are a number of reasons why such partnerships make sense for public gardens. First, to be accessible to an increasingly diverse population, many programs need to be based where people reside or gather, rather than at the gardens themselves. Also, addressing environmental hazards in our poorest neighborhoods and connecting people with plants supports the priority that many public gardens now place on environmental stewardship. Through collaborations, gardens can be invaluable to initiatives that involve urban greening, while their partners contribute social capital and other organizational, financial, or logistical aspects. Public gardens can learn from organizations that have already entered into partnerships that resulted in successful initiatives. Our book provides multiple examples of partnerships that gardens have formed with school districts, municipal governments, community foundations, businesses, and neighborhood associations. Some of those partnerships are described below. In each example, partners have brought their particular strengths to the project, with the result that together they have had a far greater impact than any could have produced on their own. Community impact is possible across a variety of sectors, including public safety, food security, educational quality, and economic development: Promoting Neighborhood Safety and Well-Being Many underserved neighborhoods are still feeling the effects of the federally supported redlining initiated in the 1930s. Unequal access to mortgages, inadequate park development, and absence of street tree plantings are all manifestations of how discriminatory housing policies still impact low income and communities of color. To provide all communities within their borough with opportunities to reverse these trends, Brooklyn Botanic Garden created the Greenest Block in Brooklyn contest in partnership with multiple local block and civic associations, the Brooklyn Borough President's office, and engaged funders. The residents who plant and beautify their individual streets are not only improving the environmental quality of their surroundings, they are also building social ties, enabling them to better address neighborhood concerns related to crime prevention, economic development, and other quality of life issues. The prizes awarded to the \\\"greenest blocks\\\" also serve as points of pride for residents, encouraging them to sustain their beautification efforts in subsequent years. Access to Healthy Foods and Promoting Healthy Lives Underinvestment also strikes poor urban communities in the paucity of food choices. Neighborhoods that lack access to a full range of grocery items, especially fresh fruits and vegetables, have been termed \\\"food deserts\\\" and are often associated with high rates of obesity, high blood pressure, and diabetes, health issues that take a toll on our medical system. To address the negative impacts of food deserts throughout the Bronx, the New York Botanical Garden (NYBG) created Bronx Green-Up, a program that annually supports up to 75 community gardens and urban farms throughout the borough. While NYBG provides horticultural expertise and planting supplies to each growing site, the program's continued success depends on a range of local nonprofit and educational partners. Among these are Farm School NYC, NYC Parks Green Thumb, Butterfly Project NYC, and the Bronx Land Trust. Through its hands-on engagement with communities, NYBG, which was once seen as an elite institution, now feels like a welcoming space for many more of its neighbors. Training and Employment Programs Throughout our metropolitan regions, prisons are overcrowded, and sentencing is often long delayed. Many incarcerated individuals are likely to return to their past behavior, resulting in a continued cycle of recidivism. Some public gardens in partnership with other community institutions have created initiatives to help break this cycle, especially for non-violent offenders. Programs such as Roots to Re-Entry (R2R), administered by the Pennsylvania Horticultural Society, treat young incarcerated men as individuals, rather than statistics. The R2R program depends on partnerships with the Bureau of Prisons, the District Attorney's Office, the Defender's Association of Philadelphia, and a web of local landscaping businesses and foundations. This synergy of participants allows the R2R program to train participants in horticultural and mechanical skills, and then helps to place them in green industry positions. Moving these young men into paying positions is enhanced by mentors who assist them as they re-enter the world. As a result, the recidivism rates for R2R participants is 30 percent compared to 65 percent for the general prison population in Philadelphia. Improving the Quality of Science Education Science education in U.S. public schools suffers from too little time devoted to the subject, a shortage of trained science teachers, and traditional curricula that focus on memorization rather than engaging students in interactive instruction. In many urban school districts, a high proportion of students also suffer from the negative impacts of poverty. These shortcomings come as the nation deals with the consequences of global climate change, loss of biodiversity, and polluted environments. The Chicago Botanic Garden (CBG) is addressing these challenges through their multi-stage Science Career Continuum. The initial stage, Science First, is an intensive, inquiry-based summer program designed to introduce middle schoolers to basic biological principles and applications. At the Garden, students are led by a team of public school science teachers and CBG staff in scientific inquiry, investigations, and observations. Those who excel are invited to participate in the College First program for eleventh and twelfth-grade that features a summer-long intensive immersion program at the Garden, including an environmental science practicum, an internship with a CBG scientist, and a monthly session with CBG staff mentors during the school year. Chicago Botanic Garden partners with a range of educational and social services organizations in the execution of these programs, including Northwestern University, City Colleges of Chicago, Chicago Scholars, Chicago Public Schools, and Hive Chicago, a network of nonprofits dedicated to connected learning. The impact of this continuum of programs is stunning: 100 percent of College First participants have graduated from high school, and 94 percent matriculated to two- or four-year colleges. These hardly exhaust the examples\u2014or the possibilities. The partnerships mentioned in this article and others described in more depth in Public Gardens and Livable Cities, will not, by themselves, solve all of the challenges of our cities. But such efforts are making a difference in creating cities that more livable for all residents. They succeed because they involve the collective expertise of both the gardens and their partners; they address real issues as identified by community members; and they reach out broadly to engage partners from across the municipal, not-for-profit, and business worlds. To learn about additional exciting collaborations between public gardens and their partners , visit our companion website at https:\/\/blogs.cornell.edu\/pglc."},{"type":"arnoldia","title":"Viburnum bracteatum 'Crimson Gem'","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25812","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070a36b.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"10","authors":"Enzenbacher, Tiffany","start_page":"50","end_page":"53","article_content":"'Crimson Gem' bracted viburnum (Viburnum bracteatum) is the first cultivar release from the Arnold Arboretum of Harvard University's new plant introduction program, Arnold Selects. This bracted viburnum's one-of-a-flower was discovered over ten years ago by the Edwards Lab (then of Brown University, now of Yale) while conducting research in the Arboretum's collections, and noted by Keeper of the Living Collections, Michael Dosmann. Reading his May 2010 database field check note, one can still feel Dosmann's excitement: \\\"inflorescences have red centers!\\\" Indeed, blossoms have a rare sterile red floret in the center, resembling the herbaceous biennial Queen Anne's lace (Daucus carota). The red floret is perched above the ivory ones, literally taking center stage. These unique flat-topped cymes are held on erect pubescent, 2-inch stems that are a whopping 1 \u00bd to 3 inches across. In fact, the red floret can be enjoyed even before the shrub comes into full flower, while its pearl-colored neighbors are still in bud. But the real show starts when the enchanting inflorescences open in their entirety in late May to early June, showering the shrub, and revealing the 'gem' in the center of practically every single inflorescence. Foliage is also appealing: emerging leaves have an ephemeral reddish hue, notably visible along the margins and between veins. Leaves are 2 to 4 inches long and 1 \u00bc to 2 \u00bd inches wide, and change to dark green as the season progresses. They are ovate with acuminate tips and serrate margins. Giving the plant its common name are the four persistent, bract-like stipules at each node (junction of leaf petiole and stem), as well as the two bracts below each inflorescence. Paired with leaf colors ranging bronze to maroon, the clusters of shiny oval to rounded fruits (drupes) are blue-black at maturity, providing autumn interest. 'Crimson Gem' was originally acquired as part of the Arboretum's commitment to plant conservation. The Arboretum is a participating member of the Center for Plant Conservation (CPC), and has holdings of ten current indigenous North American species on the CPC threatened plant list. Founded in 1984 at the Arboretum, and with the national headquarters currently at the San Diego Zoo Safari Park in Escondido, California, CPC is unique partnership of conservationist organizations, such as botanical gardens and arboreta. Its aim is to preserve threatened North American plant taxa by ex situ conservation focused on acquisition and species stewardship. Bracted viburnum is considered critically imperiled (global conservation status G1) and is endangered in the wild\u2014mainly due to limestone quarrying, but also by clearing and logging operations. Only a few populations remain in the southeastern states of Alabama, Georgia, and Tennessee. In its current range, it is found selectively as an understory shrub growing in open deciduous woodlands and along rivers, particularly on the ledges of the Coosa River, which begins in Rome, Georgia before entering Alabama, and on the escarpment of the Cumberland Plateau (southern part of the Appalachian Mountain Plateau). With CPC collections at the forefront of Arboretum institutional priorities in the mid-1980s, assistant plant propagator Rob Nicholson embarked on multiple expeditions to facilitate ex situ conservation. \\\"I basically would go to the herbarium, compile a list of locales for the targeted species, and off I went,\\\" Nicholson recalls. \\\"Once in awhile I would connect with a botanist who knew a species and where to find it \u2026 but it was mostly solo hunting.\\\" (Nicholson was a prolific collector for the Arnold and other institutions; see his account of the expedition to collect Pinus krempfii beginning on page 32.) Nicholson's investigations led him to travel to Tennessee and Georgia in October 1987, where scarce native populations of bracted viburnum are present. Twenty-one acquisitions from the very full weeklong Southeast State Expedition were accessioned. The trip was characterized by \\\"long 16-hour days \u2026 and long miles.\\\" Other taxa of interest on the trip included Fraser fir (Abies fraseri), endangered due to the balsam wooly adelgid; southern bush honeysuckle (Diervilla sessilifolia), considered threatened in its native state of Tennessee; and longstalk holly (Ilex collina), vulnerable with only 4,000 individuals in the wild. With so many target taxa acquired, Nicholson's CPC expedition was a success, but not to the extent that he could ever have imagined. Nicholson found the 'Crimson Gem' parent plant growing in Franklin County, Tennessee. Flowering in late spring, this bracted viburnum's precious bloom could not have been observed when Nicholson made the collection on October 10th. Though it was in its fall fruiting phase, he recalls that no fruit was present, so he harvested a division\u2014the sole means of preserving this crucial imperiled taxon. To hedge his bet, Nicholson took three divisions, two of which remain in the collections. Three days later, Nicholson visited Floyd County, Georgia, and collected other bracted viburnum divisions on the bluffs overlooking the Coosa River. These are also planted in the living collections, but don't share their relatives' unusual attribute. Bracted viburnum divisions were cataloged upon receipt at the Arboretum, and 'Crimson Gem' was given the accession number of 1067-87. Plants were dutifully nurtured at the Dana Greenhouses at the Arboretum, and then aptly planted at the Viburnum Collection\/Greenhouse border (likely so Nicholson could keep tabs on his valuable acquisition accomplishment) in 1992. In 2005, stem cuttings were taken of this endangered (and extraordinary) plant to increase the Arboretum's ex situ holdings. The resulting clone, with the same ruby \\\"gem,\\\" was planted in the Leventritt Shrub and Vine Garden. With over thirty years in the Arboretum landscape, 'Crimson Gem' has stood the test of time. It has provided outstanding seasonal interest, and has been reliable year after year in a shrub border. It would also be well utilized if planted as a hedge, or in mass. 'Crimson Gem' is adaptable to different soil conditions\u2014it has certainly tolerated the Arboretum's acidic soils just fine, though it is native to the alkaline soil of limestone woods. 'Crimson Gem' cuttings were just provided to nursery partners, so it will be several years before this new cultivar is available in the trade. However, it will no doubt be the jewel of gardens someday, and we can all have that magical \\\"aha\\\" moment\u2014flowers have red centers!"},{"type":"arnoldia","title":"The Timber Culture Act","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25813","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070a76e.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"9","authors":"Elkin, Rosetta S.","start_page":"40","end_page":"49","article_content":"The Timber Culture Act (1873) was a radical means to secure resources in otherwise treeless environments. Its stated claim provided entry \\\"for the cultivation of timber which are prairie lands or other lands devoid of timber.\\\"1 Its passing mobilized the protocols and promises of affidavit to deed land across the prairie states. The act was followed by land entries and land patents, the final steps in securing land that had already been peopled for thousands of years by hundreds of Native American tribes.2 Nebraska\u2014a prairie state\u2014took the lead in land entry numbers, tallies, and acres, which frames the itinerary of tree planting as a form of land tenure predicated on afforestation because Nebraska is principally a dry prairie grassland.3 Therefore, the Timber Culture Act sanctioned the substitution and deletion of dryland ecology, in this case both the extant prairie biome and the skillful cultivation of its inhabitants. A quick review of the associated institutional language makes evident the use of authority, hinged on administrative tasks. It further implicates the individual farmer as a developer, invested in the act of swapping thick grassy cover for individual tree units in order to establish an enterprise. Where 160 acres are taken, at least five must be plowed within one year from date of entry. The following, or second year, said five acres must be actually cultivated to crops or otherwise, and another five acres must be plowed. The third year the first five acres must be planted to trees, tree seeds or cuttings, and the second five acres actually cultivated to crop or otherwise. The fourth year, the second five acres must be planted to trees, tree seeds or cuttings, making, at the end of the fourth year, ten acres thus planted to trees.4 A \\\"tree claim\\\" could be filled in by anyone, and in return no more than 160 acres was acquired, with the specification that 40 acres of the claim must be tree planted, an effective means to ensure settlement. According to one claimant, \\\"the section of land specified in my said application is composed exclusively of prairie lands, or other lands devoid of timber; that this filing and entry is made for the cultivation of timber and that I have made said application in good faith.\\\"5 Units coalesced local and regional affairs, advancing federal and legislative ambitions. But the cost was only refined in the fine print, which specified that no fewer than 2,700 tree units could be planted on any ten-acre block.6 It was a staggering number that led to copious tallies, monitoring programs, and the need for reportage. Each territorial procedure (survey, reportage) reinforced the hopelessness of the task: water was scarce, winds were strong, erosion seemed erratic, and the quantity of units could not be sustained. To plow and maintain 160 acres with a mix of crops and trees was a challenge that took hold for very few claims.7 Managing 675 living trees on each acre was akin to owning a product, and growing trees was the best way to hold onto your land and avoid the cancelation of your claim. The number of entries made under the Timber Culture Act records area in acres, although territory could only be claimed if individual tree units were calculated and submitted for approval. This kind of accounting leaves out more than it includes, marking both the beginning of afforestation- linked landscape management and the rise of incentive-based policy maladapted to drylands. In Nebraska alone almost nine million acres were claimed under the act, but final proof was only made for trees planted across two million acres.8 Since alterations to the land were less predictable than imagined, the gradual accumulation of expertise was necessary to further instill confidence in afforestation. The newly formed federal government certainly did not want western expansion to be deemed a failure. Not surprisingly, this tension between success and failure required expertise, which arrived in the form of technical manuals, suitable species lists, and machinery types: the index of expertise. In the radical ecology of afforestation projects, the concept of failure is critical to the vocabulary of management, whereby catastrophe becomes a claim made by a specialist rather than a circumstance of ill-aligned biotic and climatic association. From a different angle, this means that expertise reinforces ecological projections through a series of calculated procedures rather than distinguishes, decodes, and evolves through direct experience with plant life. In fact, the plant really had nothing to do with it. Thus, failures in tallying units set the stage for the assimilation of expertise, which was equally an opportunity for the emerging business of arboriculture to become the science of forestry. The Timber Culture Act mobilized substantial takings across drylands, revealing that trees were valued as an important natural resource before forestry emerged as a discipline. At the time, forestry was a colonial economy, a means to import and spread viable European plants across the American continent and send New World novelties back to Europe.9 The botanical studies and the natural sciences were far too remote from the actual efforts of farming, explicated by the disparity created between increasing knowledge of plant parts in botany and the transmission of their application into common use, through horticulture. The more practical \\\"cultures\\\"\u2014 including the planting of trees in cities, crops for a homestead, or flowers for public display\u2014were the domain of arboriculture, horticulture, or agriculture, for instance. Botanical studies were firmly indifferent to practical endeavors in much the same ways as farmers or ranchers are indifferent to scientific study. Botany's elevated status ensured that it did not labor, and its scientific status protected it from ever having to do so. So long as these two practices (botany and horticulture) remain entirely distinct and in organized separation, a summarized reading of plant life is multiplied into the space of physical, earthly matters. Imagine a world of individual farmers working across vastly differentiated territories. Each effort represents a struggle to hold boundaries across a landscape without trees. Consequently, the failure to establish trees compelled professionalization. Failure was pinned on individual farmers and expertly manipulated as a means to assert more control over territory.10 It was not attributed to the ill calibration of species, quantity or the insistence of inserting trees into a vast, treeless landscape. Rather, lands beyond the hundredth meridian look empty from the shady groves of the East Coast. Timber and crops were needed to support a growing population, and the prairies represented a blank canvas. As a result, tree culture across the Plains was predicated on the need for protection from the harsh climatic conditions, which offered a correspondingly reduced description of the grassland biome.11 In order to increase agricultural production by sheltering farmlands from strong winds, each belt, break, or wall represented more than local shelter, firewood, or fuel: it was an opportunity to nationalize timber. The replicable model of planting a shelterbelt catalyzes the farmer as an agent of the federal economy by suppressing indigenous practices and local acquaintance. The language of this authority eliminates conjecture through levels of survey, reportage, and authoritative lists that impress with calculation. Of the total area (Great Plains region), some 56 percent lends itself to shelterbelt planting, about 39 percent is difficult to plant, and 5 percent is entirely unfit for planting.12 According to this federal explanation, there is nothing essential, intermediary, or material to constrain the accumulation of units. This description instrumentalized efforts to liberate the vacant spaces between meridians. Settlement necessitated clearing and cutting forest to make space for cultivation, as the character of cheap nature ultimately supported expansion. Both tree planting and tree felling were exploitative regimes that consolidated control across the American continent. As we have seen, the difference was contextual; trees were cleared to make space for cultivation along the arborized East Coast, and trees were planted as pioneers moved west into the treeless prairies to cultivate land. In both cases, trees with clear human advantage were selected, while plants that grew of their own accord, or spontaneously, were overlooked and rarely stabilized in the procedures of index. It took decades of energy, planting, and felling before allegiance to the spontaneous plants was described by Charles Sprague Sargent: Many years ago, when I first realized the difficulty of obtaining any true knowledge of the trees in this country, I formed the plan of writing a Silva which should contain an account of all the species that grow spontaneously in the forests of North America.13 The study of trees as spontaneous organisms with indigenous characteristics was described in The Silva of North America (1891), a work of botanical inquiry that detailed 412 species, with reference to growth habit linked to illustrated maps that indicated distribution. 14 Most significantly, Sargent, the director of the At the time, Sargent held the position of professor of arboriculture, which ensured that his tenure at the arboretum would be focused on applied engagement with the landscape, with \\\"plants in a living state.\\\" Sargent valued the associations between the individual tree, and its necessary relationship with soil, climate, and human codependency. His treatise provided a careful account of spontaneous woody plants, distinguishing between rainfall patterns and elevation, factors that limit tree growth at the hundredth meridian. This is the boundary of aridity that reveals the prairies as dryland. The variable edge is endowed with a watercolor- like transparency that emulates this bond between climate and plant life. Armed with the notes on \\\"wood producing capacity,\\\" Sargent's treatise would go on to become the first technical forestry report of the newly created Division of Forestry in the Department of Agriculture in 1886. The Silva of North America elevates claims in drylands, as forestry reaches into scientific study to nurture its advance. Of note is that while the profession of forestry is confirmed in the United States in the European tradition, it expands to include planting in order to secure extractable resources. The varied institutional settings between botany as a science and forestry as a profession entangle the study of plant life in the discrepancies and difficulties reports, as forestry read gaps in the maps and botanists read their extents. Rather than carefully consider the findings of Sargent's botanical fieldwork, remote structures deployed the literal blank space of his mapping efforts as a means to recontextualize the prairies as a tree-planting endeavor, converting biomes and replacing thick, fibrous rhizomes with woody plant units. The appropriations of forestry transformed Silva into an operations manual\u2014or a masterplan\u2014which begins to explain how vast quantities of grassland expanse and ancient reserves of pastoral acreage become the jurisdiction of federal foresters. The map was \\\"empty,\\\" a curse of drylands the world over, but it was an opportunity for the industrious. The insertion of trees infused meaning into New World expansion in the prairie states, as demand for ownership lines and boundaries enabled land claims hinged on tree planting. Afforestation emerges as forestry (not botany) and disseminates plant life as a \\\"tree unit,\\\" transferring knowledge of resource management to each small-hold farmer. In this way, particularly profitable and predictable plant species are endorsed over others, superimposing units on extant plant life. New World Forest Problems 1891 published. The Silva of North America (Sargent) 1901 renamed. Division of Forestry becomes the Bureau of Forestry 1905 renamed. Bureau of Forestry becomes Forest Service 1910 published. The Fight for Conservation (Pinchot) In the American consciousness, the origins of environmentalism are tied to the ethics of conservation, first recognized as forest conservation. The lineage of conservation, like many historic accounts, is entirely dependent on the narrator. For instance, ecological thought in America was first affirmed as plant ecology, the study of plant life in the environment. What is worth bearing in mind is that despite its significance, plant ecology did not proceed as the science of consequence in the grassland biome. Rather, forestry administered and studied the prairie formation. The reasons for this odd fragmentation of knowledge are slippery to pin down, but it is worth noting the role of Gifford Pinchot (1865-1946). Pinchot was more of a politician than a forester, and he acted as the first head of the newly established Forest Service in 1905. Under his leadership, forestry progressed from a division within the Department of Agriculture (USDA) to a separate bureau responsible for forest policy across the whole land surface of the United States. The Forest Service emerged\u2014not coincidentally\u2014 as the nation demanded unprecedented quantities of timber for fuel and construction. Charged with a simplified problem statement, Pinchot pushed forest conservation into unchartered territory. The influence of George Perkins Marsh (1801-82) motivated Pinchot's interest in the debates of conservation. In Man and Nature (1864), Marsh determined that humans were a major geologic influence, famously asserting that \\\"man is everywhere a disturbing agent,\\\" a landmark statement in both ecological and conservation studies.16 Marsh was neither forester nor plant ecologist but articulated an unparalleled understanding of the role of plant life in preserving the landscape, in how a plant's connection to the soil maintains the substrate and prevents or reduces erosion. According to biographer David Lowenthal, the success of Marsh's rhetoric is found in his caution concerning impending disaster, by addressing what he termed \\\"New World\\\" forest problems. Marsh challenged the general belief that human impact on nature was generally negligible, explicating a global tendency toward wasted natural resources. Armed with Man and Nature, Pinchot adopts conservation as forest policy. In a sticky exchange of authority, conservation and forestry coalesce as both ethic and science, alternatively sanctioning felling and planting trees regardless of context. Despite Marsh's warning, the Forest Service appropriated conservation as a \\\"solution\\\" to forest devastation, seizing upon Marsh's warning of old-world forest decline as a motivation to enforce national policy. Conservation policy progressed through extant forests and forest plantations, as plant ecology evolved as a scientific discipline in the prairie states. Remarkably, the difference was not correlated, although ecology revealed the role of plants in forming the territory and conservation uncovered the role of human agency over the territory. Rather, conservation was annexed to the profession of forestry, authorizing the Forest Service to oversee the planting of grasslands, prairies, agricultural lots, deserts, urban centers, and high mountains, concealing their extractive procedures in the outlines of conservation. Tree-planting projects begin to materialize in the millions of acres as the soil is planted, replanted, aplanted, seeded, fenced, and labeled. The landscape is objectified, appearing and disappearing through measurement and calculation. In this way, felling and planting develop synonymously, rendering the environmental messages of Man and Nature were translated as generalized statements with gross ecological miscalculations. For instance, the notion that tree planting would bring rain to the arid West was appropriated to encourage settlers to move to ever-dryer lands.17 The assumed property that followed tree planting is a clever disguise for afforestation. As Lowenthal's biography verifies, Marsh's message was pillaged for such \\\"proof\\\" that rainfall and prosperity followed tree planting. This nexus between national forest policy and conservationist thought is repeatedly stressed in Pinchot's The Fight for Conservation (1910): \\\"As a forester I am glad to believe that conservation began with forestry, and that the principles which govern the Forest Service in particular and forestry in general are also the ideas that control conservation.\\\"18 The fight for conservation was taken up by the Forest Service, exploiting the growing concern over forest devastation in order to plant and thus control more federal land. Tree cover emerges in the lexicon of conservation. Land, absent \\\"cover,\\\" is manipulated into land that can potentially receive cover. Tree units add up, when acres of \\\"cover\\\" are procedurally converted by the practices of forestry, as tree planting is sanctioned in non-treed environments. Consider the following passage. The crown has more to do with the life of the tree than its other parts, for the most important processes in the reproduction of the tree and the digestion of its food take place in the crown. For this reason, and because we can control its shape and size more easily and directly than that of the roots or trunk, the crown is of special interest to the forester. It is almost exclusively with the crowns that he has to deal in tending a crop of trees and preparing the way for succeeding generations. As they stand together in the forest, the crowns of trees form a broken shelter, which is usually spoken of as the leaf canopy, but which may be better called the cover.19 Land, absent \\\"cover,\\\" represents millions of acres of potential federal territory. The conversion of \\\"crown\\\" or \\\"leaf canopy\\\" into \\\"cover\\\" had a number of spatial consequences that will be addressed further. For now, the consequences are especially poignant because the root system is set aside in the first pages. Regulators, policy makers, and the forest service administrators intentionally reduced the growing plant to standing timber. The consequence is that the federal government had no use for the nascent scholarship of grassland science or the complications of plant ecology. The hidden intelligence of rhizomatic formation could not reaffirm exploitation or capital return, one of the most fundamental reasons why canopy continues to prevail in carbon-capture calculations. This is what professional expertise does when it lays claim to fact and asserts the power of disqualification instead. Pinchot's use of the term cover not only left out the aliveness of plants; it delineated the absence or presence of trees. The social implications and the negligence toward practical knowledge or public intelligence were already a form of expert persuasion upon which the profession of forestry is established: treeless lands can be conserved by planting trees. This raises another question: What are the spatial\u2014 and thus social\u2014consequences of such abridged problem statements? If Marsh explicated the \\\"problems\\\" of human action, Pinchot clarified that forestry had solutions, a defense strategy that he claims as a \\\"virile evolution of the campaign for conservation of the nation's resources.\\\"20 Conveniently, the solution relies on the interrelated goals of national expansion and resource extraction. By associating tree planting with conservation, Pinchot appealed to President Teddy Roosevelt, who pioneered land conservation armed with Pinchot's treatises.21 Thus, a fragmented translation of Man and Nature is interpreted by statistical proof in Pinchot's The Fight for Conservation. At the time of Man and Nature's publication, federal regulations had already secured the first Forestry Service in the federal Department of Agriculture, established a forestry program at Yale University, and set aside more than 150 national forests as reserve areas. Such policy and regulatory potency have lasting spatial repercussions. Particular to the success of forestry was the reduction of national geography to manageable units, as conservation became a capital project.22 Thus, the terms of conservation mitigated the affiliation between planting and felling, but conservation was fortified by yield management, the precision of index. This is significant to the ways in which management and development continue to prosper through the invention of crisis. Specification in Units Afforestation hides in the manipulation of specifications, loudly advocating for tree planting while quietly converting biomes: cut here, fill there or fell here, plant there. The cyclical arrangement is wedged between industrial and ecological intentions, necessitating expertise in both analyzing extant lands and constructing entirely new ones. Not surprisingly, the interdependencies between forest and field mobilize the need for more and more units as a primary measure of converting biomes, or planting trees to secure timber. The forester does not deal in individual trees destined for a purpose but on masses of tree units, because the individual tree has value only as part of the whole. At the same time that Weaver was excavating earthy workshops and painstakingly detailing the depth and intricacy of the prairie formation, the character of American expansionism was emerging in tree units, as domesticated plants balanced settlement. The accumulation and distribution of homogenous units triumphed over the diverse, interconnected, spontaneous, and multiple, preventing any expression from unscientific worlds and erasing the intimate aspects of daily life where social, cultural, environmental, and plant life mingle. Armed with an antidote to treelessness, forest policy worked its way across 170 million acres of American dryland by stabilizing a sanctioned list of tree varieties, despite the warnings that cycles that transform forest to field to plow flatten diversity and destroy soil by field wash at both an unprecedented scale and an unprecedented speed due to plow and policy. The Timber Culture Act confirmed that only the most reliable, predictable plants could be confidently inserted, so these updated lists confirmed only trees that could persist in prairie soil. To facilitate planting, the specification of tree units was linear, and in the American context catalyzed the shelterbelt typology. Dryland shelterbelts are additive, unlike the shelterbelts in Europe or those found in the American East, which are primarily composed of remnant forest. Such belts are the residues of clearing, burning, or grading procedures. The plants that remain intact are often called hedgerows or windbreaks, and are registered vertically through a prolific, enduring accumulation of biota that remain deep in the rhizosphere. In other words, the trees, shrubs, and herbs of the former landscape are unbroken, and continue to thrive in diversity and habitat. As a network of accretive forces and biological activity, they are extremely effective at slowing erosive forces. The subtractive force of clearing land produces an increase in mounds and layers within the hedgerows, as branches and rocks are thrown into piles and endure. Each mature specimen also maintains vigorous seed banks and robust sprouting stumps. When a plant is inserted as an individual organism, copied and pasted in neat rows, it has limited ability to produce relations in the soil. This is due to the constitution of the rhizosphere, in which a process between microorganisms, fungus, and soil biota is stratified in cooperation with other neighboring organisms. Plants, left behind with their major root systems and mycorrhizal relationships intact, continue to provide reliable growth patterns. The connection of each remnant to the soil advances the entire network, an interdependent and persistent spatial structure, which pulses through the soil. In contrast, shelterbelt protocols in drylands insert plant units into flattened, cleared, turned, and exposed soil. Plants\u2014treated as objects\u2014are forced to instigate each belowground relationship anew before any achievement can be registered aboveground. A shelterbelt not only has behaviors, domains, and unknowns; it has a spatial geometry that often necessitates two landowners, as it straddles property lines. Further, this geometric configuration not only extends a thickened line and extrudes a height gradient; it extends the root zone in both the horizontal and vertical dimensions. The roots require irrigation, nutrition, and time in order to assert aboveground effects. As a result, local disputes escalate along with the behavior of any trees that display significant mobility, as some plants spread more quickly than others. While a shelterbelt is designed to grow in height and girth, it is certainly not expected to migrate or move. If a shelterbelt is \\\"successful,\\\" it is only because the plant evolved persistent root morphology and reproductive capacity, a feature that cannot be rendered visible or valuated economically. Yet conflict arises if the plant is successful enough to be found encroaching beyond property lines, or into productive farmland.23 For instance, the behavior of some woody roots will outcompete annual crops for resources such as moisture and nutrients, which makes them a nuisance to farmers. In shelterbelt culture, trees are deemed beneficial only if they provide undeviating protection, and are deemed a menace if they overcome specifications. The correlation between additive and subtracted spatial geometry illuminates the friction between developers and their environment during this period. Tree planting in the Plains exemplified this struggle, as failure to adapt was decidedly unacceptable and un-American.24 Resolve only increases in the space between dying and trying, a feature of most developers bent on return. But it was the lack of consensus between investments that finally led to the appeal for government support.25 Just as the American pioneers resisted treelessness, they could not cooperate on investment between property limits. Ownership is a personal discipline, as the contested fence and neighbor relationship suitably explicates. In a largescale planted system, these thresholds prove mutually beneficial. Yet the owner who initially broke ground, transplanted, or seeded the rows tended to be the one to water, manage, and maintain it, despite shared outcomes. Effort was the developer's investment, and resistance has the power to affect the system it endorses. Simultaneous to the decades when professional forestry, sedentary agriculture, and public acquisition were determining the American landscape, conservation collapsed neatly into agencies, services, institutes, and organizations. Political agendas merged with development strategies through the protection or abuse of natural resources, so that access and recreation became the most salient features of the vistas, lakes, peaks, and other wonders of natural beauty.26 Forestry lurks in the background as the landscape becomes a cultural and social construct, devoid of individual life-forms beyond the human. The symmetrical competition between growth and yield, or materializing and subtracting, currently calculates carbon credits through additive operations and timber volume in deductive increments. At once, local populations are led to believe that \\\"billion tree\\\" campaigns and \\\"great green walls\\\" are pronouncing progress, but growth statistics and depletion metrics are isolated from accountability and context. These flip-side tactics benefit from quick rotations that leverage tree planting as a means to elevate control through yield, in a world troubled by ongoing crisis. When dealing in units, the authority of forestation selectively neglects the less visible attributes of plant life, including individual plant behavior, symbiotic relationships, and the concealed roots and rhizomes that form and deform the soil within which biomes are produced. Because of aboveground visibility, development budgets, and technical encouragement, projects that eagerly plant trees under the rubric of afforestation continue to garner international funds and approval despite the haunting conspiracy of replacing biomes and commercializing plant life. The prowess of progress that celebrates biome conversion creates as it destroys, a central tenet of afforestation. rosetta s. elkin is associate professor and academic director of landscape architecture at Pratt Institute, principal of Practice Landscape, and research associate at the Arnold Arboretum. endnotes: 1. The full Timber Culture Act is available in Thomas Donaldson, The Public Domain: Its History, with Statistics (Washington, D.C.: Government Printing Office, 1884), 1093. 2. Roxanne Dunbar-Ortiz, An Indigenous Peoples' History of the United States (Boston: Beacon, 2014). 3. For more on Nebraska as a \\\"tree-planting state,\\\" see John F. Freeman, \\\"Trees for High Plains,\\\" in High Plains Horticulture: A History (Boulder: University Press of Colorado, 2008), 19-32. 4. Donaldson, The Public Domain, 1092. 5. Donaldson, 1094; Timber Culture Act, application October 3, 1878, Individual Claimant, NB. 6. Benjamin H. Hibbard, A History of the Public Land Policies (Madison: University of Wisconsin Press, 1965), 422. 7. Accounts of individual claims are covered in relation to the landscape in Charles Barron McIntosh, The Nebraska Sand Hills: The Human Landscape (Lincoln: University of Nebraska Press, 1996). 8. Everett Newfon Dick, Conquering the Great American Desert: Nebraska (Lincoln: Nebraska State Historical Society, 1975). 9. Philip J. Pauly, Fruits and Plains: The Horticultural Transformation of America (Cambridge, Mass.: Harvard University Press, 2007). 10. C. B. McIntosh, \\\"Use and Abuse of the Timber Culture Act,\\\" Annals of the Association of American Geographers 65, no. 3 (1975): 347-62. 11. In a description of the region, an introductory study by Raphael Zon of the Forest Service explains that \\\"the climatic conditions become less favorable for plant growth from east to west.\\\" Lake States Forest Experiment Station, U.S. Forest Service, Possibilities of Shelterbelt Planting in the Plains Region (Washington, D.C.: Government Printing Office, 1935), 3. 12. Lake States Forest Experiment Station, 7. 13. Charles Sprague Sargent, The Silva of North America: A Description of the Trees Which Grow Naturally in North America Exclusive of Mexico (Boston: Houghton, Mifflin, 1891), v. 14. See also Charles Sprague Sargent, Sixteen Maps Accompanying Report on Forest Trees of North America (Washington, D.C.: U.S. Census Office, 1884). 15. Sargent, The Silva of North America, v. 16. George P. Marsh, Man and Nature (New York: Charles Scribner, 1864), 14. 17. On the embrace of Pinchot's policies by Americans, see Michael Williams, Americans and Their Forests: A Historical Geography (Cambridge: Cambridge University Press, 1989), 393-411. 18. Gifford Pinchot, The Fight for Conservation (New York: Doubleday, Page, 1910). 19. Gifford Pinchot, A Primer of Forestry (Washington: Government Printing Office, 1903), 7. 20. Pinchot, The Fight for Conservation, iv. 21. David Lowenthal, George Perkins Marsh, Prophet of Conservation, Weyerhaeuser Environmental Book (Seattle: University of Washington Press, 2000), 9. 22. Lowenthal, 10-12. 23. Office of Information, U.S. Forest Service, Prairie-Plains Region Shelterbelt Project (Washington, D.C.: U.S. Forest Service, 1946). 24. Wilmon H. Droze, Trees, Prairies, and People (Denton: Texas Woman's University, 1977), 30. 25. Droze, 22. 26. There is tremendous literature on the history of conservation. See, for instance, George Perkins Marsh, Man and Nature, ed. David Lowenthal (Seattle: University of Washington Press, 1864); and Douglas Helms and Susan L. Flader, eds., The History of Soil and Water Co"},{"type":"arnoldia","title":"Two Poems","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25814","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070ab26.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"8","authors":"Fjeld, Jessica","start_page":"39","end_page":"39","article_content":"Early Early I descend the stairs like a child's newly-acquired marionette No part in predictable relation to the others In the kitchen on the top shelf is a cardboard bankers box Full of boiling water Good The lilac's leaves all withered but for months it has had a hundred green little buds, firm little pregnancies at the tip of each branch I am watching them daily to gauge their intentions I think you should find it surprising that given the time we spend on our feet, it's so difficult to learn to stand on our hands How the ground presses up The cement is compromised, full of hope and sand I tell the children it's a bunker due to the bunk beds They want to show the beds to visitors but instead we teach them moral rectitude, screaming, subsisting on sugar and edible species of fungus Their teeth sink into cake like gravity sinks into the snow laying thick on the mountainside The mutinous avalanche turns the trees upside down but the trees are prepared Their roots and branches mirrored in all relevant respects jessica fjeld is the author of Redwork (2018) and the chapbooks The Tide (2010) and On animate life (2006), for which she received the Poetry Society of America's Chapbook Fellowship. Also an attorney and academic, Fjeld is the Assistant Director of the Harvard Law School Cyberlaw Clinic at the Berkman Klein Center for Internet & Society, where she focuses on supporting the work of creatives, archivists, and human rights defenders as that work intersects with emerging technology. She lives with her family in Somerville, MA."},{"type":"arnoldia","title":"Pinus krempfii Does Battle","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25815","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070ab6a.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"7","authors":"Nicholson, Rob","start_page":"32","end_page":"38","article_content":"All the plants of a given country are at war one with another. The first who establish themselves by chance in a particular spot tend, by the mere occupancy of space, to exclude other species\u2014the greater choke the smaller, the longest-lived replace those which last for a shorter period, the more prolific gradually make themselves masters of the ground, which species multiplying more slowly would otherwise fill. \u2014Augustin Pyramus de Candolle 1820 Writing more than a decade before Charles Darwin embarked on the Beagle, Swiss botanist Augustin Pyramus de Candolle (1778-1841) foreshadowed the idea of the survival of the fittest when he wrote of the ecological competition between plants, a competition with inevitable winners and losers. Over geological history, the groups of plants that cloak the earth have shifted dramatically with changes in climate, and as new groups of plants evolved and entered the battle for territorial supremacy. Between the Devonian and early Cretaceous periods (about 380 to 125 million years before present), the world's flora saw the rise and dominance of gymnosperms: conifers, cycads, ginkgos, and others. In the early Cretaceous, angiosperms, or flowering plants, began to evolve, proliferating into the hundreds of families, now some ninety percent of all plant species, which now dominate vast areas of the planet. Gymnosperms (roughly one percent of extant plant species) surrendered huge expanses of land and were pushed to high altitudes, the boreal region, or into cohabitation with the new \\\"masters of the ground,\\\" flowering plants. Modern ecologists shun the vivid language of \\\"the war between the plants,\\\" couching this battle in terms like \\\"sequential clade competition\\\" and the \\\"active displacement hypothesis.\\\" But with global warming, the shrinking of the boreal forest and mountaintop refuges will only accelerate the gymnosperms' loss of territory. This largely human-induced change may be more rapid than any yet withstood by the group. Ebb and Flow The largest family in the gymnosperms, Pinaceae, the Pine family, includes 232 species, with 119 described as pines in the genus Pinus. Of these true pines, the IUCN lists nineteen as falling in the most dire categories: vulnerable, endangered, or critically endangered. Pinus has one of the widest natural ranges of any genus of trees on the planet, both in terms of latitudinal distribution and altitudinal distribution, and is also one of the most widely planted of trees. Predominantly native to the Northern Hemisphere, pines grow from 72 degrees North latitude in Siberia to just below the equator in the mountains of Sumatra, and range in elevation from coastal species like Pinus rigida and Pinus banksiana to pines in Mexico and the Himalayas growing well above 10,000 feet elevation. Still one of the most poorly known pines, Pinus krempfii of Vietnam is also highly unusual. Its broad, flat leaves can measure 12 mm wide over a length of 10 cm, unlike the more familiar thin needles of species like white pine, Pinus strobus. In the tropical forests it inhabits, it grows to be a colossal tree, towering over the surrounding flora. The species was described in 1921 by the French botanist M. Krempf, in the Nha Trang vicinity, where it is no longer found. Never widely collected, its specimens in cultivation number in single digits outside of Vietnam. A tropical holdout in the war of the plants, P. krempfii isn't faring as well in the competition for collection. Prior to the discovery of Krempf's pine, paleobotanists had recorded finds of fossils of a broad-needled Pinaceae. The current climatic conditions where Pinus krempfii is found are warm, humid, and equitable. These conditions prevailed during the Paleocene Epoch (56-66 MYA), during which time a 'boreotropical flora' dominated much of the middle latitudes. Gymnosperms such as Glyptostrobus, Sequoia and Taxodium were components of this flora, and, since this epoch, their ranges have been severely contracted. This would have been a period in which P. krempfii or its precursors may have had a far wider range. A diminution of favorable conditions for the boreotropical flora during the Eocene (34-56 MYA), coupled with increasing competition from angiosperms, may have severely reduced the potential range for a conifer adapted to mild conditions. Two fossil discoveries in 2021 in Northern Thailand and Yunnan relate to P. krempfii, and show it once had a greater range. Like Metasequoia glyptostroboides (dawn redwood), it now exists in a highly restricted area. This perhaps adds P. krempfii to the list of \\\"living fossils,\\\" such as dawn redwood and Wollemi pine\u2014plants that were known as fossils before they were linked to extant populations. Dr. Tim Brodribb, a former Fellow at the Harvard Forest, has visited the wild stands of Pinus krempfii, and his 2008 report with Dr. Taylor Field is to my mind the best recent ecological paper. He argues that Krempf's pine behaves more like a tropical gymnosperm Podocarpus, noting that the tree \\\"shows photosynthetic, hydraulic and anatomical characteristics more akin to a southern hemisphere podocarp than a pine tree,\\\" with leaves that show a \\\"striking convergence with the flattened leaves of rainforest Podocarpaceae\\\" as well. But what really catches my eye is Brodribb's profoundly de Candollean view of the species. Krempf's pine, he notes, offers the first evidence that \\\"Pinus has the physiological (and anatomical) capabilities to invade equatorial evergreen forest. The question remains whether the evolution of broad leaves and shade tolerance is a recent development, representing a new southward invasion of the genus, or alternatively the remnant of an ancient and unsuccessful invasion.\\\" Whether recent or ancient, invasion is a constant. Collecting for the Smith College Conservatory in the autumn of 1998, I teamed with Dr. Shu-Miaw Chaw of Taipei's Academy of Sciences to collect rare conifers of Taiwan and Vietnam. We wished to bring into cultivation many rare and endangered species for future research purposes, and Dr. Chaw was to collect DNA samples of conifers for her work on the evolution and phylogeny of gymnosperms. Our research team was to have an extraordinarily successful trip, collecting such rarities as Amentotaxus formosana, Cephalotaxus wilsoniana, Cephalotaxus mannii, Calocedrus macrolepis, Taxus wallichiana, Keteleeria evelyniana, Podocarpus nerifolius\u2014and one of our primary goals, Pinus krempfii. Once in Hanoi, we began our journey with a courtesy call to Dr. Le Thi Xuan, the head of the Biotechnology Institute, whom we knew from prior collecting trips for Taxus\/Taxol research. She had been instrumental in linking us to Dr. Tran Ngoc Ninh, a capable field botanist and expert in the Rubiaceae. As time was tight due to our early flight, we only had time to maneuver quickly through local alleyways to her home and take tea. Our old friend was ill, struggled to move and speak, but insisted on extending hospitality to us. We socialized and exchanged gifts and then briskly headed to the airport for the flight south. As we were leaving, she pulled me aside. \\\"Be careful in the \u0110\u00e0 La\u0323t Forest,\\\" she whispered. \\\"It is full of danger.\\\" \\\"What kind of danger?\\\" I replied. \\\"Two-legged or four-legged?\\\" \\\"Danger of every kind,\\\" she replied, in a grace note worthy of Joseph Conrad. After two flights from Hanoi, we landed at \u0110\u00e0 La\u0323t airport and drove upward into the hills, where the city lay, at 4900 feet (1500 m), home to over 100,000 people. A balmy summer capital for many of the regimes that have ruled Vietnam, \u0110\u00e0 La\u0323 t boasts a miniature Eiffel Tower as a souvenir of occupation by the French, along with the fine French bread and French roast coffee that can be found in the cafes. At the southern terminus of the Annamite range of mountains, one of the southernmost pieces of high ground on the Asian landmass, a number of conifer genera reach the southernmost extent of their ranges. Isolated here for millions of years, they may possess unusual genetic characteristics. We stayed at the Biological Institute, located in a former Catholic monastery on the town's outskirts. It was built by Catholic monks in 1950 but lost to the Communist takeover in 1954. Despite the apparent low budget, the institute had a tissue culture lab and was involved in conserving the many endemic plant and animal species of the region. In the small nursery outside grew a number of conifer species we sought, including Taxus wallichiana, Dacrydium elatum, Fokenia hodginsii, and Podocarpus neriifolius. The former Director of the Arnold Arboretum, Dr. Peter Ashton, who had been to \u0110\u00e0 La\u0323 t, had informed me that a specimen of Pinus krempfii also grew nearby. In C\u00f4ng Ty Park on the outskirts of town, we found a Keteleeria evelyniana in cone, a member of the pine family, which grows well in our southern states but is poorly known horticulturally. No more than fifty feet from it, we found our prize: a specimen of Krempf's pine. Looking like a broad-leafed Podocarpus from only ten feet away, though on close examination it held its new \\\"candles\\\" like a pine, and smelled like a pine when crushed. We happily collected needles and herbarium sheets. The next day we drove to Xuan Tho, a section of \u0110\u00e0 La\u0323 t, to see a stand of native Taxus wallichiana, the Himalayan yew, and to collect cuttings for the Taxus collection at Smith College, a collection used extensively for anti-cancer research. It was the height of Vietnam's coffee boom, and the whole area showed a recent conversion to plantations, as Viet coffee gained in stature among gourmets. At 1420 meters, we walked down a slope toward a small stream bed. Nepenthes smilesii, a vinelike pitcher plant, was growing among the grassy understory of this pine forest. Only two yews remained, one a substantial specimen, eighty feet high with a basal diameter of four to five feet. On our return to the institute, we stopped at the home of a local plant collector, a forester, in whose lathe house grew numerous orchids alongside a locally-collected specimen of Cephalotaxus mannii, a gymnosperm within the Cephalotaxaceae, from which he generously provided us cuttings. After a hard night's rain, the dawn broke clear, and we loaded our war-vintage Russian jeep and headed out of town, driving past schoolchildren trudging down the road in neat blue uniforms. We wove through a quilt of grassy pinelands and carefully tended, productive family farms. Nine kilometers from the institute, we halted beside a roadside grove of Keteleeria evelyniana that had recently suffered through a forest fire. Plants from the collection we made there currently grow with Dr. Jason Smith of Florida State University. Another species, Keteleeria roulettii, had been reported as endemic to the \u0110\u00e0 La\u0323t area, based on the observation of longer needles. But from our observations of K. evelyniana, it was clear that fire had caused a series of new shoots to regenerate longer juvenile foliage, and this character was probably the basis of a false \\\"new\\\" species. It is no longer recognized. We continued on the muddy red-orange road upward through a pine savannah, very similar to the longleaf pine forests of southern Georgia, into an unpopulated and relatively undisturbed piece of geography. About twenty-eight kilometers from the institute, we stopped at 1700 meters altitude in the area known as L\u00e1n Chanh. Putting our packs to our backs, we marched to the crest of a small hill. Looking down into the valley, we could see a drastic change in the vegetation: the piney grassland became a primeval evergreen forest, moist, dark, and lush. Rising above it all, a towering example of the pine we had come so far to see and collect, Pinus krempfii. Rather than having a spire-like habit, this forest giant had a broad crown, more like an oak than any pine I could remember seeing. We eased downslope toward the sounds of birds trilling and a river's flow, crossing a transition zone so narrow we felt as though we had stepped through a glass wall into a terrarium. Suddenly we trod upon a soft spongy humus soil dotted with ferns and seedlings of the evergreen trees around us. So many primitive angiosperms were present\u2014Magnolia of the Magnoliaceae, Illicium of the Illiciaceae, Sacandra of the Chloranthaceae\u2014it seemed a forest undisturbed since the Cretaceous era. Orchids were common, having fallen from the branches above, species of Dendrobium, Bulbophyllum and Epigeneium that would be precious to any orchid fancier. We made our way to the large pine we had seen from the hilltop, which proved a tree of mythic proportions: about 120 feet high with a trunk circumference of eighteen feet, the first branch at a height of seventy-five feet. Climbing the tree to harvest cones would have been unreasonably risky without climbing equipment, so we had to forage for what we could find on the ground. Our party of five took to combing the forest floor, looking for fallen cones from prior years that might still harbor a seed or two, or for seedlings and saplings we might dig up. At one point I searched farther away from the party only to be cautioned by Dr. Tran to come back into the fold. \\\"There are tigers,\\\" he warned. We found 4 younger trees and about twenty seedlings to bring to our respective institutes. The leaves were so unlike any I had ever seen, curving blades 4 mm wide by 6 cm long, held in pairs. Cones that we found in the forest duff were easily recognized as pine. We also collected cuttings of Podocarpus neriifolius, along with some of the epiphytic orchids fallen from their perches above, and a smattering of ferns and seed. Our Vietnamese colleagues seemed intent on getting out of the forest as quickly as possible, and counseled against crossing the valley to look for more trees. I recollected Dr. Xuan's warning in Hanoi, and was relieved to think that we had avoided any peril beyond the usual rigors of the field. In any case, I already had collected a cache of rarities for our Smith College conservatory, and we had precious DNA samples for genetic work. We rose out of the ravine, over the grassy slopes and down to our waiting vehicle. We lunched at the side of the road, fine French-style baguettes, local fruit and imported soft cheese. Our success fueled our good mood and we sat on the side of the road, talking about our various plant specialties and favorite species. I glanced down at my feet and saw in the road's gutter a weathered bullet casing, American made, a probable relict from the war that once tore both nations apart. It was my final collection of the day. The few seedlings we brought out seemed to languish in the heat of our New England summer, and eventually died. Despite numerous hormone treatments, none of our cuttings rooted (though this is typical for pines). Other than the lonely plant at the Cong Ty Park in \u0110\u00e0 La\u0323 t, the only plant I could find in cultivation is held in the conservatory at Royal Botanic Garden Edinburgh. Groves of Krempf's pine also are found within the boundaries of the Bidoup Nui Ba National Park, established in 2004, though their remoteness may make them susceptible to illegal timber poaching. Pinus krempfii remains aloof, distant and beckoning still. Given its great height, future collectors may need collection equipment like that used for Sequoiadendron: a bow and stringed arrow followed by ascension gear, with skilled arborists acquiring a supply of seed for ex-situ work. Growing ex-situ specimens outside native ranges requires a combination of factors: the correct climate conditions (whether in a conservatory or outside), a long-term infrastructure of skilled horticulturists, and institutional will. The correct climate conditions would seem to be the biggest stumbling block for Krempf's pine, as it grows in montane tropical or cloud-forest conditions without extremes of cold or heat. Its hardiness is not known. Some of the mildest areas of the California coast and upper altitudes in Puerto Rico may serve as potential US sites, though hurricanes would be a factor in Puerto Rico. But as was shown with Metasequoia, it can be a fool's game to make hardiness assumptions based on a current restricted population range. As elsewhere, the war of the trees continues in Vietnam, though thankfully the wars of the humans there have subsided. For all its rarity, Pinus krempfii is a magnificent holdout against angiosperm domination, one of the most amazing species I have ever seen in a long career collecting plants. In discussing local politics, Dr. Xuan had once told me, \\\"In Vietnam, nothing is possible but everything is possible\\\"; the latter part of that truism seems to apply to the flora as well."},{"type":"arnoldia","title":"The Wild Hydrangeas of Mt. Cuba Center","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25816","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070af6d.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"6","authors":"Hoadley, Sam","start_page":"20","end_page":"31","article_content":"When I bought my house five years ago in Chester County, Pennsylvania, I inherited a landscape with a garden tradition dating back to the 1940s. Mature specimens of choice woody plants are bountiful and include a grove of mature dawn redwoods (Metasequoia glyptostroboides), three massive franklinia (Franklinia alatamaha), and one large and aged smooth hydrangea (Hydrangea arborescens). The identity of this smooth hydrangea cultivar is not known, but due to the age and the substantial sprawling nature of the planting I suspect it to be Hydrangea arborescens 'Grandiflora', formerly sold under the name 'Hills of Snow'. This cultivar was originally selected from a wild population of Hydrangea arborescens in the early twentieth century, and is still available from specialty growers today. Hydrangea arborescens 'Grandiflora,' along with the similar and more popular H. arborescens 'Annabelle', are staples of American horticulture that are readily recognized by expert and novice gardeners alike. Their oversized, dome-shaped inflorescences borne in June and July have classic ornamental appeal, but these cultivars are not without their shortcomings. Both 'Grandiflora' and 'Annabelle' have floppy habits, a flaw that is exacerbated by summer storms right when the blooms are at their peak. For this reason, I had always considered these cultivars to be horticultural relics and, admittedly, gave little consideration to the use of the species in modern landscapes. My insular outlook was quickly challenged, however, thanks to my inheritance of a comprehensive trial of smooth hydrangea when I was hired for my current role of Manager of Horticultural Research at Mt. Cuba Center in Hockessin, Delaware, in March of 2019. Mt. Cuba Center, situated in the rolling hills of New Castle County, Delaware, is the former estate of Pamela and Lammot du Pont Copeland. The Copelands envisioned that Mt. Cuba could be a place to inspire an appreciation for the beauty and value of native plants and spark a commitment to protect the habitats that sustain them. This original intention lives on as Mt. Cuba Center's core mission, and continues to guide the organization to this day. After Mrs. Copeland passed away in 2001, the transition from private estate to public garden began in earnest. In 2002, the Copelands' cut-flower plot was repurposed to serve as a trial garden. In this space, native species and cultivars were grown side by side to determine their horticultural qualities in a low-maintenance common garden setting. The first research report was published in 2005, describing findings from a threeyear trial on asters, including floral display, habit, cultural adaptability, winter hardiness in Zone 7a, and disease resistance. This publication highlighted and recommended the top-performing plants to gardeners and nurseries in the mid-Atlantic region, and served as a template for future research reports. A report on Echinacea was released in 2009, and in 2011, the trial garden was redesigned with a new perimeter fence, brick walkways, and a shade structure. The shade structure proved to be an invaluable tool, not just for trialing shade-loving species, but also for comparing the performance and adaptability of trialed plants when they are cultivated in both sun and shade. In 2013, Mt. Cuba Center opened its doors to regular public visitation for the first time, and the trial garden welcomed visitors to observe and interact with trials in progress. Under the deft management of George Coombs, publication of the trial garden's research reports became a highly anticipated resource valued by the public and nursery industry. Trials were established in a staggered rotation so that four trials run concurrently, with one trial concluding each year. In addition to promoting plants with excellent garden qualities, the Coreopsis, Monarda, and Phlox reports aspired to address the ecological services that these species and cultivars provide by collecting comparative pollinator data on the various species and cultivars in each trial. Pollinator studies quickly became an important component of Mt. Cuba's trial program, and have allowed consumers to make informed decisions about which plants have the most capacity for attracting and supporting pollinators in their home landscapes Quantitative and qualitative horticultural data are collected in the trial garden on a weekly basis between April and September. Scores are assigned that reflect habit, form, foliage, and other ornamental qualities that the plant may exhibit. For example, a weak plant with a floppy habit might receive a low score, while a vigorous plant with a robust and nonfloppy habit would receive a high score. A separate rating is assigned to the floral display when the plants are in bloom, and an additional rating assesses disease resistance, which becomes especially important when observing plants that have a known susceptibility to specific pathogens. At the end of each season an overall score is generated for each plant. At the conclusion of the three-to-five-year evaluation, the scores from each year are averaged to determine the final horticultural score. In the most recent trials of hydrangea and Echinacea, supplemental points were then awarded to plants that attracted the most pollinators. This supports our initiative of promoting plants that are both beautiful and also have wildlife value. The top performing plants are then featured in subsequent research reports. For some gardeners, selecting a native plant for its ornamental qualities is enough, but for others this is only part of the greater story of providing habitat and food sources for wildlife in their home landscape. Pollinator data are collected by a cadre of Mt. Cuba Center's citizen scientists, called the Pollinator Watch Team. On a near-daily basis, these volunteers observe plants in bloom for 60 seconds and collect information on insect visitors. Counts are made on one plant per accession, or single inflorescences if blooms are numerous or plants are prohibitively large. The number of pollinator visits is recorded, along with the time, weather data, and additional observations. In some trials, all pollinators that visit the plant or inflorescence are counted, while in others, only targeted groups such as butterflies or hummingbirds are recorded. The results of these studies help homeowners make informed decisions about which plants in our evaluations have the most capacity to attract and support insect pollinators. These data become particularly relevant as natural habitats face increasing threat of destruction. Home gardens are uniquely poised to temper those losses, and can help provide small oases for wildlife in the midst of food deserts comprised of the turfgrass and non-native foundation plantings that make up the majority of home and commercial landscapes today. By pointing people in the direction of beautiful and beneficial native plants, Mt. Cuba Center can help people become conservators. When I was hired as Manager of Horticultural Research in the early spring of 2019, I was greeted with four in-progress trials: Helenium, Echinacea (the second trial of this genus), smooth hydrangea (or \\\"wild\\\" hydrangea, as they are called at Mt. Cuba Center), and Carex. Smooth or \\\"wild\\\" hydrangea represented the first woody plant trial at Mt. Cuba and had just begun the third year of a five-year evaluation. While the study of woody plant material represented a step into new territory for the trial garden, it was a logical one. There is an increasing demand from homeowners for reliable information on alternatives to non-native woody landscape plants that are low maintenance, beautiful, and providers of wildlife value whenever possible. Included in the smooth hydrangea evaluation were three species of hydrangea native to the eastern United States: Hydrangea arborescens, Hydrangea cinerea, and Hydrangea radiata. Hydrangea arborescens was the primary species evaluated, and is the wild version of most of the twenty-six cultivars in the trial. Hydrangea arborescens, or smooth hydrangea, has the largest native range of the three species and can be found in much of the eastern and central United States, often on shaded moist slopes. Hydrangea cinerea, or ashy hydrangea, is exceedingly rare in cultivation and can be found primarily in the central and southeastern United States. The best identifying feature of ashy hydrangea is the concentration of white to grey pubescence on the stems and the backs of their leaves, which differs from the relatively hairless foliage of Hydrangea arborescens. Hydrangea radiata, or silver-leaf hydrangea, has the smallest native range compared to Hydrangea arborescens and Hydrangea cinerea and can only be found in a handful of states in the southern Appalachian Mountains. The most striking feature of Hydrangea radiata is the bright white undersides of the leaves making this perhaps the most easily identified and inherently ornamental species of the three in our trial. Silver-leaf hydrangea was also the first to bloom in our trial in mid- June while the other two species started blooming in early July each year. While the three species vary slightly in size, habit, and overall appearance, each flowers on new wood and produces superficially similar inflorescences. The three species, including two distinct accessions of Hydrangea radiata, were all propagated from wild collected stock from the collection at Mt. Cuba Center, and these plants acted as the controls against which all of the cultivars would be compared. Twenty-five cultivars were then sourced from various arboreta and nurseries, in most cases representing material available to homeowners at that time. All twenty-nine hydrangea accessions were grown in full sun and nineteen cultivars were also grown in shade for comparison. Since these shrubs bloom on new wood and flower buds are not produced until the start of the growing season, smooth hydrangea can be heavily pruned in late spring and flowers are still reliably produced in June and July. To assess response to pruning, one example of each hydrangea grown in full sun was cut back to approximately six to eight inches from the ground in late March 2019, 2020, and 2021. The remaining examples of each hydrangea served as the control group and were left unpruned for the duration of the evaluation. That first spring at Mt. Cuba, before the hydrangea had even leafed out, I was struck by the diversity of form, texture, and even stem color. Subtle variations between similar cultivars and species were readily apparent when the plants were grown side by side. It became obvious to me how short-sighted I had been when I had written off smooth hydrangea for my own garden use. There was clearly so much more to this group of plants than the old standbys, like Hydrangea arborescens 'Grandiflora' and 'Annabelle'. As spring progressed and data collection began, I looked forward to what I knew would be a spectacular floral display in June and July. When the time came, the flowers were indeed breathtaking, and included a concert of white and pink inflorescences that fell into two distinct categories: lacecaps and mopheads. Lacecap smooth hydrangea species and cultivars produce flat-topped, corymbose inflorescences that are primarily composed of masses of fertile flowers that number anywhere from 800 to 2,000 individual flowers. Around the perimeter of many lacecap flowerheads are a small number of sterile flowers that give the inflorescences their trademark lacy appearance. Lacecap inflorescences are the predominant flower form found in wild populations of Hydrangea arborescens, and offer visiting pollinators highly accessible nectar and pollen rewards. Hydrangea arborescens and Hydrangea radiata both produced lacecap inflorescences that averaged around 4.5 inches in diameter, although the sterile flowers of the silverleaf hydrangea were noticeably larger than those of the smooth hydrangeas. The fertile flowers of Hydrangea radiata were also less tightly clustered than those of Hydrangea arborescens. Hydrangea cinerea, on the other hand, produced the smallest diameter flower heads in the trial at around 3.5 inches on average, and also some of the smallest sterile flowers. Despite the smaller diameter, the inflorescences of the ashy hydrangeas were tightly packed with fertile flowers and were heavily visited by pollinators. In contrast, mophead hydrangea blooms are often more dome-shaped and produce a much higher number of sterile flowers per inflorescence, giving them their iconic billowy cloud-like appearance. In most cases, fertile flowers are still produced, but in much lower numbers than their lacecap counterparts. In addition, these fertile flowers are often sequestered within the inflorescences, reducing their accessibility for some insects. Some mophead smooth hydrangeas such as Hydrangea arborescens 'Annabelle' are selections made from naturally occurring anomalies found in nature. However, not all mophead smooth hydrangeas in cultivation are the products of simple selection. Wild-origin mutations form the genetic building blocks utilized by modern plant breeders to cross and further select smooth hydrangeas. The progeny of these breeding programs feature ornamental and horticultural improvements as well as novel garden traits. The trial at Mt. Cuba was planted with mophead cultivars concentrated on one end of the planting bed and lacecap cultivars and species planted on the other. As I walked through the blooming mophead cultivars, the inflorescences were beautiful, but the surroundings were quiet and relatively devoid of insect pollinators. As I continued down the path to the lacecap hydrangeas, this changed, and I was greeted with an immersive audible and visual experience composed of hundreds if not thousands of insects. Bumblebees, wasps, beetles, true bugs, and flies busily collected pollen and nectar from the fertile flowers of the lacecap inflorescences in a frenzy of activity. I was in awe, not just because of the huge number of insects I was witnessing, but also by the stark contrast between the two flower forms and the obvious preference that the various pollinators displayed for the lacecap smooth hydrangeas. One lacecap cultivar in particular, Hydrangea arborescens 'Haas' Halo', completely captivated me. Not only was it covered with pollinators but here was a robust plant with an incredible floral display that rivaled any of the mophead cultivars in the trial. This was an example of a plant that encapsulated Mt. Cuba's mission. This ornamentally superior and garden-worthy native plant inspired an appreciation for its beauty and for the ecological value that this plant was clearly providing to scores of insect pollinators. I was so taken by Hydrangea arborescens 'Haas' Halo' that I ordered one for my home garden that same day. This single experience in the trial garden constituted a personal paradigm shift in the way I looked at horticulture and my own gardening practice. There was room for beauty and ornamental interest, which had always been a priority, but ecological value did not have to be sacrificed. In fact, gardening for insects and other wildlife quickly became a focus for me, and has been a deciding factor for the vast majority of my home-garden additions ever since. This one plant has inspired me to take conservation action by planting a native shrub with high ecological value in my home landscape. And I was not alone. Over the next three years, I fielded countless questions about Hydrangea arborescens 'Haas' Halo' from visitors who had experienced this plant for the first time in the trial garden just as I had. They all wanted to know where they could buy one. This is exactly the kind of inspiration and conservation action that I imagine the Copelands envisioned when they first conceived of the idea to one day welcome the public into their home and gardens. In 2021, the five-year trial of smooth hydrangea was complete and the results were in. The top-performing hydrangeas were identified, and pollinator counts were tallied. The pollinator data confirmed what could be readily observed when the plants were in bloom each year. Insects overwhelmingly preferred lacecap hydrangeas over mopheads. That is not to say that all mopheads are completely devoid of pollinator value. In fact, some attracted a reasonable number of insects. The mophead that attracted the most insects in the trial was Hydrangea arborescens 'NCHA2' (Invincibelle\u00ae Spirit II) which, maybe not surprisingly, produced a greater proportion of accessible fertile flowers than other mopheads. While some compromises exist where you can have both the mophead aesthetic as well as some pollinator value no mophead hydrangea received enough pollinator visits to be directly compared to any of the lacecaps. Interestingly, the hydrangea with the lowest number of pollinator visits was Hydrangea arborescens 'Hayes Starburst'. This cultivar exhibits mophead inflorescences but is an extreme case of genetic variation within the species. While most mophead smooth hydrangeas produce a reduced number of fertile flowers, Hydrangea arborescens 'Haye's Starburst' produced no observable fertile flowers. No fertile flowers mean no pollen or nectar and therefore no pollinator benefits. This lack of pollinator value was directly reflected in the low pollinator count. Interestingly, there were three exceptions that broke the rule that lacecaps are better than mopheads for pollinators: Hydrangea arborescens 'Riven Lace', Hydrangea arborescens 'Emerald Lace', and Hydrangea arborescens 'Green Dragon'. These are relatively compact plants with lacecap inflorescences and distinctive dissected foliage. These three cultivars are so similar that it is widely believed that they are actually the same, genetically identical plant that was named and introduced into the horticultural trade on three separate occasions. Their inability to attract insect pollinators remains a mystery, but could be attributed to several factors, including the possibility that these cultivars do not offer the same quality or abundance of nectar or pollen as other lacecaps hydrangeas in the trial. From a purely ornamental perspective, most of the nineteen smooth hydrangea cultivars grown in shade performed better than the specimens in full sun. The primary reason for this is that shade-grown plants avoided issues such as foliar and floral burn as well as premature defoliation. Inflorescences of the hydrangeas grown in shade retained their form for months after the fertile flowers had finished blooming in July, and aged from white to attractive shades of lime green. Hydrangea radiata and its cultivars struggled the most when grown in full sun, particularly during the heat of the summer. These were always the first to show signs of heat and drought stress and were the first to start dropping leaves. This species is best reserved for use in woodland edges and shade plantings in the mid-Atlantic region. At Mt. Cuba Center, Hydrangea radiata is grown to perfection in the shady naturalistic gardens where the silvery undersides of the leaves add movement and interest to the landscape and their lacecap inflorescences attract insect pollinators in droves. To my surprise, a handful of pink flowering hydrangeas performed exceptionally well in full sun, even better than in shade, including Hydrangea arborescens 'NCHA2' (Invincibelle\u00ae Spirit II) and Hydrangea arborescens 'NCHA4' (Incrediball\u00ae Blush). These two cultivars can be successfully sited in full sun locations (six or more hours of sun a day) in the region surrounding Mt. Cuba Center if they are planted in moisture-retentive but well-drained soils. Good soil moisture remains an essential component for successful cultivation of any of the smooth hydrangeas planted in full sun or near full sun conditions. The cutback comparison conducted over three seasons revealed some interesting trends. There was a slight delay in bloom (generally one to two weeks) in cutback plants and fewer but larger inflorescences. This increased diameter of the inflorescences can be attributed to a return to a more vigorous juvenile state in the cutback shrubs. On average the inflorescence increase was around thirty percent, while some examples increased between fifty to eighty percent in diameter. For some plants this effect was particularly noticeable especially if the inflorescences produced on a non-cutback plant were already of substantial size. For example, the cut back version of Hydrangea arborescens 'Haas' Halo', which produced the largest inflorescences (8\\\") of the trial on the control plants, produced a dramatic display of flower heads that were nearly a foot across. Cutback hydrangeas also formed a more compact habit in many cases. Some exceptions to this rule were compact cultivars such as Hydrangea arborescens 'NCHA5' (Invincibelle\u00ae Wee White) and Hydrangea arborescens 'NCHA3' (Invincibelle\u00ae Ruby) where there were few discernable differences between cutback and control plants. One unexpected effect of the cutback study was that cutback plants had improved sun tolerance. Not only might the cutback plants have had more efficient water-transporting stems, but their established root systems were supporting fewer water-demanding leaves compared to the controls. However, I would not recommend annual heavy pruning to increase sun tolerance over the long run or to make larger plants permanently more accessible to smaller planting spaces. The best practice is to select the right cultivar for the right garden location based on the ultimate size of the unpruned plants and the goals of the gardener. I recommend a more restrained approach to pruning, where only a handful of the oldest stems are removed on a yearly basis, to maintain some juvenile vigor in your hydrangeas. One unexpected but welcome benefit of hydrangea pruning in the trial garden was the observation of square-headed wasps (Crabronidae) using the pithy stems as a nesting site. On several occasions we observed adult wasps excavating the spongy, pith-filled cores of recently pruned stems. Once the cavity had been established, the adult wasp could be seen placing paralyzed flies in the hollowed-out stems for its larvae to feed on. The chambers were then sealed, protecting the developing larvae within. The larva eventually pupate, and emerge the following spring as adult wasps. Perhaps this observation would inspire people to tolerate and even embrace some dead wood in their shrubs if they knew what a benefit they could potentially be to wildlife. However, if spring pruning is required, it is still feasible to prune out full stems even if they have been colonized without interrupting wasp's life cycle. The discarded stems can be placed intact somewhere to allow emergence to occur later that year. In addition, there is an opportunity to further utilize the pruned stems by simply cutting them to lengths of eight inches to a foot and the bundling the stems together. These makeshift bee nesting sites will only further amplify the bee and wasp habitat in your landscape by using raw materials provided by your hydrangeas. In addition, Hydrangea arborescens is the larval host plant for the hydrangea sphinx moth and the hydrangea leaftier moth, although neither species was observed in the trial. Nine cultivars of smooth hydrangea were identified as top performers, and the highest rated plant of all was indeed Hydrangea arborescens 'Haas' Halo'. This cultivar, besides being my personal favorite smooth hydrangea, offers the perfect combination of horticultural merit and ecological value. This exemplary form of smooth hydrangea was originally selected by Frederick Ray in 2008, plantsman and former Delaware Valley College horticulture professor, from the Pennsylvania garden of Joan Haas. It was chosen for its upright, vigorous growth, and for its lacecap flower heads, upwards of eight inches in diameter, that display a greater number of sterile flowers around the perimeter than those of typical Hydrangea arborescens. A comparable, but decidedly more obscure, selection called Hydrangea arborescens 'Mary Nell' made the top performer list as well. It too produces abnormally large lacecap inflorescences, distinguished by a double ring of sterile flowers around the perimeter of each flower head. Hydrangea arborescens 'Mary Nell' was selected and named by Joseph McDaniel, who also introduced Hydrangea arborescens 'Annabelle' in the 1960s. Unfortunately, Hydrangea arborescens 'Mary Nell' is rarely commercially available despite its substantial garden and wildlife value. Two more lacecap cultivars made the top performer list. Hydrangea arborescens 'Dardom' (White Dome\u00ae) is an older cultivar originally selected in Belgium in 1997 and introduced by Proven Winners\u00ae in the early 2000s. It boasts large lacecap inflorescences and attracted the highest numbers of pollinators of any hydrangea in the trial. Hydrangea arborescens 'Total Eclipse' is a vigorous, slightly more compact, and earlier-blooming selection of the species made by Jim Pyler of Natural Landscapes Nursery in West Grove, Pennsylvania. This cultivar also attracted the third most pollinators of any hydrangea in the evaluation. The remaining five top-performing hydrangeas from the trial produce mophead inflorescences. While these plants may not have the pollinator value offered by the lacecap top performers, they are still highly recommended from an ornamental perspective for gardens of the mid-Atlantic. Hydrangea arborescens 'Abetwo' (Incrediball\u00ae) and Hydrangea arborescens 'Bounty' are two white-blooming mopheads that offer excellent alternatives to Hydrangea arborescens 'Annabelle' and Hydrangea arborescens 'Grandiflora'. Each features the classic hydrangea aesthetic that gardeners have valued for more than a century, but improve on their predecessors with sturdy stems that are resistant to flopping. Hydrangea 'SMNHALR' (Lime Rickey\u00ae), a unique mophead hydrangea from Spring Meadow Nursery, displays undeniably attractive, lime-green inflorescences in mid-June that are quickly followed by raspberry-colored fertile flowers later in the month. The sterile flowers rapidly fade to alabaster before reverting to their former green color as the fertile flowers complete their bloom. Last but not least are Hydrangea arborescens 'NCHA4' (Incrediball\u00ae Blush) and Hydrangea arborescens 'NCHA2' (Invincibelle\u00ae Spirit II). Both cultivars are some of the finest examples of the cutting-edge plant breeding that continues to push the boundaries of what is thought possible with hydrangea, redbuds, and dogwoods amongst many other woody plant genera coming from North Carolina State University and Thomas Ranney. These two cultivars feature attractive pink mophead inflorescences that are well supported by sturdy stems. Hydrangea arborescens 'NCHA4' (Incrediball\u00ae Blush) is a great option for smaller landscapes thanks to its semi-compact habit of four to five feet in height and width, while Hydrangea arborescens 'NCHA2' (Invincibelle\u00ae Spirit II) eventually forms a much larger plant reaching six feet in height and eight feet in width. For more information on the top performing smooth hydrangeas from Mt. Cuba Center's latest trial, visit mtcubacenter.org\/research\/trial-garden\/. For me, the smooth hydrangea trial represents a turning point in my personal gardening journey which, just like the garden itself, is constantly evolving and is never truly complete. I went from largely looking at plants as a decoration for my home landscape, an admittedly narrow view, to choosing plants not just for me but for wildlife that I can support and provide refuge for in my garden. Seeing the surge in insects and birds and other wildlife at my home, a trend which I would like to think directly correlates to my native plant choices, has been an incredibly rewarding experience, and is so much more than just cultivating a place of objective beauty. It provides me with a purpose, the same purpose that the Copelands had when they envisioned the future of their property that would eventually become Mt. Cuba Center. Thanks to this trial, I have a newfound appreciation for the Hydrangea arborescens 'Grandiflora' that still resides at my home in Kennett Square, although my future hydrangea selections will be made for the benefit of pollinators. My personal journey into the world on native hydrangeas has not concluded with this trial, and in fact will only be expanded upon with Mt. Cuba Center's newly planted evaluation of oakleaf hydrangeas. This trial will be unique in that, for the first time, we are trialing a single species and comparing its many cultivated forms. Hydrangea quercifolia has many seasons of interest: it is well known for the oak-shaped leaves for which it is named, white to pink panicle inflorescences in summer, incredible burgundy fall color, and exfoliating cinnamon-colored bark that can be admired during winter months. The trial will assess cultivars that are tried-and-true garden classics as well as a handful of newcomers to the horticultural market. The oakleaf hydrangea trial will be sited in full sun; however, most of the subjects will also be grown under our shade structure for comparison. This will be an interesting trial for observing pollinator preference, because the percentage of fertile flowers per inflorescence in cultivated forms and selections of this species is quite variable. Oakleaf hydrangeas are one of my all-time favorite shrubs and I know I am in for more horticultural and ecological inspiration in the trial garden at Mt. Cuba Center before the results are of this latest trial are published in 2027."},{"type":"arnoldia","title":"Helwingia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25817","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070b325.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"5","authors":"Gapinski, Andrew","start_page":"18","end_page":"19","article_content":"As a student of horticulture in the Midwest of the US, I was fascinated by the floral characteristics of the eastern redbud (Cercis canadensis). The reddish-purple flowers and brown pea-like fruits are borne directly out of the bark of older wood of main trunks and stems rather than newly emerging growth. This phenomenon, known as cauliflory, is most prevalent in tropical species including coffee and cocoa. Although still intrigued every time I see it, I am even more enthralled by epiphylly, when flowers and fruits (as well as other structures) grow attached to leaves (often the midvein). At the Arnold Arboretum, epiphyllous flowering can be observed on a single specimen of Helwingia japonica (AA# 912*MASS) in the Explorers Garden, received in 1880, likely a propagule from the first introduction of the species into European cultivation. Alone in the Helwingiaceae, genus Helwingia contains four species occurring from the Himalayas to Japan. Either evergreen or deciduous, all are shrubby, epiphyllous, and dioecious (bearing male and female flowers on separate plants). The Arnold's 1880 specimen is male, and currently lacks a paired female plant for successful reproduction. In 2018, I traveled to the Shennongjia region of Hubei Province, China, as part of the North America-China Plant Exploration Consortium. Along with important germplasm for conservation, we collected seeds of two species of Helwingia: H. japonica and H. chinensis. Both were found growing on shaded rock outcrops in rich pockets of humus soil in association with the conifer Torreya fargesii. Helwingia japonica's broad, deciduous leaves sport umbels of tiny, four-petaled flowers in spring, developing into pea-sized black drupes in autumn. Helwingia chinensis, by contrast, is a semi-evergreen species with narrow leaves bearing purplish male flowers on long stalks (pedicels) in groups of 4-5; female flowers are nearly sessile (unstalked) and in groups of 1-3 with fruits ripening to a glossy cherry red. We now have one plant from each species in the Arboretum's Dana Greenhouses. The H. chinensis plant, a male, bloomed copiously this year. Over the past 150 years, we have made several attempts to cultivate this species in the collections\u2014each to no avail. If this collection is successful, we'll need to find it a mate by collecting it again."},{"type":"arnoldia","title":"The Habitat of Childhood","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25818","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070b728.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"4","authors":"Caballero, Ana Maria","start_page":"14","end_page":"16","article_content":"Recently I took a small group of kindergartners on a forest walk atop an esker at the Arnold Arboretum. A treecovered, glacial ridge, it was a cool and shady spot on an otherwise sunny, hot, dry day. A red robin seemed to be leading the way, lowering its head while running and lifting it when stopped, keeping an eye on us as we observed it carefully. It disappeared behind a downed log covered in moss. The children were immediately drawn to it, touching the green velvet and looking for spiders and ants. Across the path, a child discovered the lower part of the downed log: a wide, hollowed-out snag that created a life size cocoon big enough for six children. They all quickly settled inside, perching on the damp decomposing wood continuing to look for spiders and other creatures. One child commented, \\\"Sometimes you can find salamanders in the woods,\\\" to which I replied that it's been so dry lately that most of them have gone deeper underground. \\\"That's because there is water underground, right?\\\" she asked. After a few silent moments she asked, \\\"Do you think we are walking on salamanders?\\\" I was so surprised by this insight and this moment: fearless children connecting to the natural world with curiosity and joy. Sharing such experiences of young children in nature, and how to promote more of them, is what prompted the convening of a panel of outdoor educators at Boston University this spring. As Outdoor Educator at the Arnold Arboretum, I support Boston Public School students who come for guided and self-guided field trips to learn in our landscape, as well as train our volunteers and offer professional development for educators throughout the year. As such, I have long followed research affirming the many benefits of nature to adults and children. Health benefits include lower blood pressure, reduced stress, a boost to the immune system, and increased anti-cancer proteins. Cognitive gains include improved focus and higher scores on standardized tests. Other benefits include fewer sleep difficulties, faster healing after illness, increased emotional resilience and stronger mental health. Recently, doctors have pegged Computer Vision Syndrome (CVS) as a direct consequence of too much screen time, and there is a documented increase in myopia (nearsightedness) in young children. Surprise! Being outdoors can alleviate these symptoms, as children look out in the distance and relax the eyes. But the benefits extend far beyond this, as I have seen with the children who attend our field study programs\u2014two-hour landscape explorations of specific life-science content, which happen in small groups led by trained volunteers. Unlike a snake-oil salesperson, I can say that we truly have the remedy for so much that ails us. I've noticed so many benefits; at the BU panel I discussed just five, which I share with you here. Active Agency in Learning Learning outdoors teaches children a sense of scale and perspective, reinforces the cyclical nature of time and space, and introduces children to experiences and words upon which later learning is built. This is how background knowledge is created. Children become the star when they experience the concept: seeing the seasons change, following the germination and growth of a plant outdoors, discovering the effects of a rainstorm on the earth, breaking apart stones, \\\"climbing\\\" the shadow of a tree limb or coming face to face with a snapping turtle for the first time. These experiences introduce vocabulary and demand that children narrate what they see, hear, and feel. In response, they use words in context, invent words creatively, and make connections between English and their first languages. This is learning that integrates physical, cognitive, emotional, and social domains; this is learning that puts the child in the driver's seat. Context Being in nature brings book and classroom learning to life by putting all that information in context. We begin with the premise that science is learning about the world around us, and the world around us is the natural world. Vocabulary and concepts make more sense when children can touch, smell, hear, and explore these same words and concepts in real life\u2014in context. It is here that we can see that a holly leaf has sharp prickles to prevent herbivory, or that a hawk does not always catch the snake, or that vines have tendrils that curl \\\"just like my hair!\\\" This story illustrates the need for context: An elementary age child confessed to one of our guides that she was afraid of squirrels, with their big front teeth, rat-looking faces, and size. When the guide asked this child \\\"how big do you think a squirrel is?\\\" the child indicated with her hand an animal about two feet tall! This child had never really seen or noticed a squirrel in context, perhaps, but only in books or movies. I have seen firsthand how being in natural context has helped children overcome perceived misconceptions and fears. Once the child catches their first bee or centipede in a bug box, all worry, fear, and disgust at the task goes away\u2014they can't get enough of digging through leaf litter looking for critters, or appreciating the pollen baskets on bees' legs. Messiness The outdoors is messy! Unexpected things happen to frustrate, delight, and surprise every day. The rain begins to fall. You come across a dead squirrel. The salamander you so very much wanted to see didn't make an appearance. Someone left their dogpoop bag on the path. A tree has to be cut down because of a storm. A coyote follows a dog on a leash. When children encounter unexpected events, they learn to manage disappointment, and perhaps fear or worry, in proportion with the encounter\u2014especially when they are involved in addressing and finding solutions for the situation. They learn to be flexible and to come up with alternatives. Tolerating discomfort that comes from unpredictable and messy nature leads children to develop a sense of personal competence. Part of growing up is learning how to release these negative TREE OF LIFE Mesozoic Mystery Our picture of the rise of the angiosperms, and the extent of plant diversity when they first arose some 140 million years ago, is far from complete. Sadly, many of the now extinct groups of non-flowering seed plants that might help reconstruct the evolutionary history of the rise of angiosperms remain undiscovered. A species of Mesozoic plant, newly described from fossil evidence in a recent issue of the American Journal of Botany, may shed light. The authors suggest possible affinities between early flowering plants and the new species, a gymnosperm, which they have named Xadzigacalix quatsinoensis (the genus name incorporates the word for \\\"plant resin\\\" [xa'dziga] in Kwak'wala, the language of the Kwakwaka'wakw nations of northwest coastal North America, on whose traditional territory the fossil was found). The case is far from resolved, however: is the plant indeed related to early angiosperms\u2014and if so, does it have anything to tell us about their origin? How are other cupulate seed-bearing groups of Mesozoic plants related to each other and to flowering plants? While such questions remain topics of palaeobotanical debate, this much is clear: the buried remnants of the Mesozoic still have many secrets yet to be revealed. Klymiuk, A. A., Rothwell, G. W., and Stockey, R. A.. 2022. A novel cupulate seed plant, Xadzigacalix quatsinoensis gen. et sp. nov., provides new insight into the Mesozoic radiation of gymnosperms. American Journal of Botany 109 (6): 966-985. Image: Crosssectional reconstruction of Xadzigacalix quatsinoensis, through apex (micropyle) of ovulate reproductive structure. Klymiuk, Rothwell, and Stockey (2022). UAPC-ALTA P15375 D. CC BY 4.0 emotions in the face of inevitable stress. If kids never figure out how to do that, they're more likely to experience severe anxiety as teenagers. These experiences outside are often the most memorable of unexpected wonders. An adult's job is to help children navigate these surprises with respect, simplicity, and empathy. Global Citizenship Children who are outdoors learn from a very young age that their actions and interactions with the natural world affect the natural world. It is one thing to be indoors and learn about trash or talk about how cutting trees is bad for the environment. It is quite another to see the effects of human activity on our world. When children see trash in the water stream, a tree branch broken under the weight of a climber, or dry, yellowing fields at the height of summer, they are primed for conversations that can center on activism and purpose. Young children who learn to love the outdoors are more likely to become adults who work to preserve it for future generations. Magic So often after a session, young children tell us that they don't want to leave, they want to live here forever, they see the Arboretum as a magical place. They are filled with possibility. Very quickly, many children come to feel a sense of belonging that comes from the peace, calm, and happiness that they experience in the outdoors. Joy. They want to share this place with their families, they want to come back. As educators, parents, and caregivers, we need to be intentional in letting children be children for as long as possible. Childhood and the natural world are wonderful spaces to be in. This habitat of childhood needs to be tended and protected. If we do it right, this magic stays with them throughout life."},{"type":"arnoldia","title":"The Sweetest Legacy","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25819","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070b76c.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"3","authors":"Reese, Carol","start_page":"11","end_page":"13","article_content":"The Holsteins were gone, and watching our hayfields and pastures being reclaimed by the wild offered a sad reminder that dairy farming was no longer economically viable in north Mississippi. The land called for new purpose, which led to years of fruit growing experimentation, searching for a crop that had, as my mother put it, \\\"maximum profit for minimal sweat equity.\\\" Eventually we found that crop in Asian persimmons (Diospyros kaki), but for years, our quest was not always fruitful (and yes, the pun is intentional). Our running joke was that our most valuable lessons were learned by killing plants, usually because they were not suited to north Mississippi. Plant death was not always the issue, however, as sometimes failure comes by other means\u2014as it did with our first project, a blueberry U-Pick venture. The local extension service had suggested rabbiteye blueberries (Vaccinium virgatum) could supply huge profits with minimal inputs. They were right about little input. On some of our ideally acid soils, those rabbiteye blueberries planted some forty years ago still produce ample fruit with minimal care. Other factors denied profitability: while U-Pick blueberries were popular, locals usually only picked a few gallons and were done for the summer. And despite our efforts to acidify those places in the field that had unsuitably high pH, large swaths of the orchard simply did not prosper. Realizing that blueberries weren't the answer, we set off on a series of jaunts to investigate other fruits and learn from other growers. We joined the North American Fruit Explorers (NAFEX) organization, attending the annual meetings that moved each year to different regions of the country. This generous community provided lifelong friends and mentors, from whom we learned to become decent grafters, with only minimal blood loss. We also learned, the hard way, that Mississippi has the perfect climate for growing disease. Plants we saw thriving in California, Michigan, Oregon, and New York swooned in our heat and humidity. My father, however, was a civil engineer, and adept at finding solutions. When it became obvious that southerners needed their own group, a drawling band of commercial and amateur growers came together to form Southern Fruit Fellowship. Like NAFEX, the meetings moved around to different regions, but always in the south, and it was in Florida that my parents bit into their first Asian persimmons (Diospyros kaki). My father said it was love at first bite. They took a picture of the smiling moment. This fruit fit the bill for a U-Pick business: it ripened in mild, beautiful October, and customers filled five-gallon buckets in a matter of minutes. The trees required little maintenance beyond pruning to develop low spreading trees, shaped to accommodate strolling fruit-pickers. Mowing between the rows was a task maintained intensively only during harvest season. There was still a learning curve. Our farm lay in Zone 7, the hardiness limit for most Asian persimmon cultivars, with wildly fluctuating temperatures in seasons of transition. We learned to plant in spring, giving trees time to settle in well before a challenging winter. We also had survival issues with trees grafted onto Diospyros lotus rootstocks, so began using American persimmon (Diospyros virginiana). They occurred naturally on our farm, so we collected and planted several seeds directly into the desired spot, selected the most vigorous of the seedlings to graft, and left a couple of ungrafted seedlings in place for a year or two alongside the newly grafted trees. These would serve as backup rootstocks, should the grafted plant come to ruin, and they helped shield the tender new graft from wind gusts or racing dogs. Later, these extras were transplanted into pots and grafted for orchard expansion or sold to local nurseries. Our first year of significant production found us in a new dilemma. Our delicious, beautiful Asian persimmons attracted very few customers! This was before the days of social media, and in an era of less cosmopolitan palates. Even more to the point, an aversion to persimmons was common among our usual clientele. As children, many had been tricked into biting into a firm American persimmon, which is astringent until it has softened. The memory of the lingering pucker caused by that unripe fruit made them understandably reluctant to be suckered (puckered?) again. Our first plantings were all non-astringent cultivars, however, and delicious even when hard as an apple. Could they be convinced to try one? Each fall, my parents set off for local farm markets and horticultural field days with cutting boards, sharp knives, and bushels of just-picked persimmons. For hours, they sliced the beautiful orange persimmons and challenged total strangers to try the crisp wedges. I helped occasionally and enjoyed seeing the expression of doubt change to delight as they gingerly bit into the crunchy sweetness. Converted! Faces went from wary to warm, and soon our booth would be buzzing with questions and conversations. Nearby Starkville had another customer base, eager and ready-made: its vibrant Asian communities. These excited new customers taught us proper pronunciation of the cultivar names, and the resulting friendships brought to our farm an unexpected and rewarding cultural exchange. Thanks to those early efforts, the crop now consistently sells out as fast as it ripens. The only advertising done is to announce opening day for picking season and hours of operation. My parents are buried not far from the orchard, and my little brother now runs the operation, but some of their early customers still come, as well as the children and grandchildren of those first customers. The tradition of picking brilliant orange fruit on a beautiful autumn day spans generations, and our farm's soil has become part of their growing families. Several years before their deaths, my parents told their seven children the only gifts they wanted were memories of time spent happily with family. I smile to think that Reese Orchard still provides that opportunity for so many, long after they are gone. It's a different measure of success, beyond an orchard's profitability, and is their enduring legacy. URBAN ECOLOGY Happy Bats, Healthy Cities COVID-19 was hard on the public image of bats, who already suffered a reputation as carriers of disease. In fact, they contribute to human health, checking populations of mosquitos and agricultural pests\u2014and many aid in pollination and seed dispersal, too. Since 2021, students supervised by Dr. Alison Robbins of Cummings School of Veterinary Medicine at Tufts University have studied bat populations in living laboratory of the Arnold Arboretum. This year, Conservation Medicine graduate student Christina DeJoseph (left, in the field) installed monitors to collect bat calls and the sounds of other species. Supported by the Arnold's Sinnott Award, DeJoseph is tracking acoustic biodiversity to map how bats contribute to public health and healthy ecosystems. With veterinary student Alex Debrindisi, DeJoseph also offered a series of public \\\"bat walks\\\" at the arboretum to foster public fascination. \\\"Taking interest in the other species we live with,\\\" DeJoseph points out, \\\"that's healthy, too.\\\" To learn more about the work of Cummings-School students in the arboretum's landscape, look for the bat \\\"wonder spots\\\" at arboretum.harvard.edu\/visit\/explore-with-us\/."},{"type":"arnoldia","title":"Incarnations of an Apple","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25820","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d070bb6f.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"2","authors":"Kaminsky, Matt","start_page":"8","end_page":"10","article_content":"It's fall in Massachusetts, and I'm traveling to a different apple orchard daily, making sure the fruit gets harvested on time from the different sites that I look after. Some days I find myself in the puckerbrush, tromping over blackberry and multiflora rose to reach piles of windfallen wild apples in abandoned pastures. Other days I spend my time with stately old trees laid out in rows. I love the variety of sites that I get to visit during this time of year. It's been very dry and mild this fall, so fruit is ripening nice and slowly. This allows plenty of time for making the rounds to the dozens of sites where I harvest. Starting with orchards located furthest south in my orbit, and ending with those furthest north, I will be busy during these twelve or so weeks ranging September to November. I've just stopped to visit to a few noteworthy, unique seedling apple trees on the side of the road, collected their offerings, and continued on. In my truck I have some ovate, crow's-egg-looking oddball apples from one tree, which tasted like pineapple, next to a crate of large, round, green, and sweet-tasting ones, and another crate of aromatic fruit from an old hollow beast just next to it, which might be an old grafted apple called Mother. But those are just the bonuses. Today, my real goal is to check on the hilltown Baldwins\u2014a very special instance of Malus domestica 'Baldwin'\u2014growing in one of my favorite places. Some years ago, I was introduced to this old orchard by a former professor of mine, located far up in the hills west of the Connecticut River, on a homestead site in Franklin County. This remote site experiences its own microclimate, a bit colder than the surrounding countryside, and the blossom and fruit development are always further behind the orchards down in the river valley. Last week, I helped the crew at an orchard seated down in the valley at the base of Mount Norwottuck cleanpick the remainder of their 'Baldwin' crop, which had been steadily dropping fruit for a couple of weeks already. I'd estimate that the 'Baldwin' orchard up in the hills hadn't dropped a single apple yet, but that they'd be coming along at their own pace, ready to harvest any day now. When I was first introduced to this special old orchard, it was in the middle of winter, and snow was falling. We couldn't tell what kind of apples the trees were. All I could see was perhaps two dozen hulking behemoths standing in the field, reveling in their dormancy. It also was apparent the trees hadn't been cared for in many, many years and badly needed pruning. My professor had described the fruit to me in rich detail, which made my imagination race with wonder and anticipation. Come the first harvest after pruning, we learned that all but just a couple of the trees were Baldwins. One of the most iconic Bay State apples, their intense fiery red skin, balanced, sharp, subacid flavor, and signature hardness (so hard it may register on the Mohs scale used by geologists) are unmistakable. However, these are not just any 'Baldwin' apples. Though they are grafted, and therefore genetically identical with all other 'Baldwin' trees, these ones hit different. They are ancient, massive trees. By counting the rings of limbs that I pruned off, and comparing to the size of the trunks, it is nearly certain that they pre-date WWII. History books and old timers alike hold that all the Baldwins in Massachusetts were killed by an unusually harsh winter in 1934, after which most growers gave up on the cultivar. Few intact 'Baldwin' orchards from that era remain. Some of the trees in this orchard have expressed the rootstock as multiple smaller trunks coming from the ground, producing different types of fruit than the main trunks, which produce Baldwin apples. Each individual tree's rootstock produces unique, intriguing apples, which don't resemble one another in any way. Their flavors are eccentric\u2014some are more interesting than the Baldwins! This signals to me that this orchard was originally grafted onto seedlings rather than a standardized rootstock. Of course, each of the seeds they planted wouldn't come true to its parent, but rather would produce a unique apple tree. This is due to the trait of extreme heterozygosity that Malus displays, hence why the auxiliary trunks produce such wildly odd fruit. Trees growing on seedling roots are generally known for their ability to thrive in adverse conditions, and indeed the soil here is somewhat inhospitable. From the fact that the main understory plant growing around the apple trees is lowbush blueberry, we can tell that it is very acidic. In most cases, soils that support prolific barrens of blueberries will prove too harsh for apple trees, which prefer a higher pH. Moreover, the extensive outcroppings of smooth, gray ledge throughout the orchard indicate that the topsoil layer is shallow. The roots have plunged through gaps and cracks in the stones to gain anchorage. It would be hard to imagine an apple more intensely flavored, saturated in color, and sound of form. Piquing, sparkling acidity is coupled with plentiful juice and lovely, lingering, winey notes of tart cherry and a full mouthfeel. The 'Baldwin' apples from these hilltown trees are smaller, firmer, redder, and more flavorful than the orchard down in the river valley, where the harvest is just finishing up. Those 'Baldwin' trees are grafted onto MM.111, a widely-established commercial rootstock common in the US since the 1950s, and are at least forty years younger than those in the hilltown orchard. The valley trees also have a leafier, more upright growing habit than the twisty, winding branches of their older counterparts. The fruit is more plentiful, larger in size, and less intense in color, with little bits of green showing through. The juicy, firm flesh is rich in flavor, with notes of cranberry, but doesn't linger and isn't as complex. They lack the intense phenolics present in the older hill Baldwins, which add texture and depth, though they're still enjoyable. The apples that I'm describing seem different from each other. In fact, if you didn't know that they were both 'Baldwin', you might not even be aware that they're the same type of apple, indeed genetically identical. How does the same scion, grafted and growing in two different locations, produce fruit so varied in appearance and flavor? An apple fruit represents the sum of all variables that are a part of the tree and its surroundings. The genetic material contained within an individual specimen is filtered through a set of environmental and cultural qualities, which can vary greatly from place to place. Nature is nurture, the marriage of genome and environment. It is subject to the influence of all life around it; the activity of other organisms interacting with the tree, the qualities of the soil it's growing in, the weather, the influence of humans, chemicals, the rootstock they are grafted on, and so on. What appearances of the fruit are caused by the interactions of insects or disease? What flavors of the fruit are imparted by a particular deficiency or nutrient surplus in the soil? What practical elements of using the fruit, such as storage quality or size, are impacted by the tree's environment? What factors are we unaware of? Once a variety like 'Baldwin' is widely established, the genetic blueprint, which is generally static and inextricable (except where mutations occur), is replicated many times via grafting. When it is grown over a wide geographic area and under cultural methods that differ greatly, we begin to see that many different incarnations of the same apple begin to appear. As with the valley Baldwins and the hilltown Baldwins, the differences, slight or significant as they may be, are immutable. When I consider that 'Baldwin', as well as most named apple varieties, originated as a seedling, it makes me wonder how different the apples I'm experiencing today might be from the original 'Baldwin' tree that was discovered in Wilmington, Massachusetts, in 1740. How did the fruit from that original 'Baldwin' tree taste? How many millions of 'Baldwin' trees have lived and died since 1740? How many incarnations of 'Baldwin' have apple growers experienced? These are the questions that run through my mind with all apples I am eating, whether some famed old heirloom like these Baldwins, or some as-yet-unknown seedling, from an old rootstock, a roadside wild apple, or a tree found growing in the woods. As seasons have come and gone, and I've had chances to taste the venerable 'Baldwin' apple from dozens of sites, from trees with different stories and different personalities, I can say with confidence that these oldest, most distinguished, and unique seedling trees produce my favorite incarnation. BOOK BRIEF Little Forest, Big Impact Mini-Forest Revolution Using the Miyawaki Method to Rapidly Rewild the World By Hannah Lewis Foreword by Paul Hawken Chelsea Green Publishing, 2022, 224 pages Protected from cutting for centuries, the forests that cluster around Japan's Shinto temples are refuges for endemic trees and the myriad creatures they support. Inspired by these biodiversity hotspots, botanist Akira Miyawaki (1928-2021) developed a method for intensive, site-specific cultivation of forest soil and cover, densely planting 30-40 regionally appropriate tree species to shelter seedlings and encourage rapid growth. As Hannah Lewis explains in Mini-Forest Revolution, the Miyawaki Method aims for big effects at hyperlocal scale, unlike most large-scale tree-planting efforts (for the fraught history and environmental impact of afforestation, read the excerpt from Rosetta Elkin's Plant Life in this issue, beginning on page 40). Author Hannah Lewis pairs case studies of Miyawaki projects from Europe to India with guidance on turning back lots into tiny forests. Editor at Biodiversity for a Living Climate, based in Cambridge, MA, Lewis first encountered the Miyawaki Method while living in France."},{"type":"arnoldia","title":"Being Green","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25821","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d0708127.jpg","year":"2022","series":null,"season":null,"volume":"79","issue_number":"3","article_sequence":"1","authors":"Dosmann, Michael S.","start_page":"1","end_page":"2","article_content":"While singing Joe Raposo's \\\"Bein' Green,\\\" Kermit the Frog laments how he blends in\u2014and fails to stand out\u2014among more colorful and flashier things. Given how many plant species (including one third of trees) are threatened with extinction, \\\"it's not that easy bein' green\\\" should be their motto, too. In urban forests, tribulations trial our street trees to the point where we measure their lifespans in years not decades, leaving cities ever hotter and far less beautiful places. Plant blindness, the term coined by James Wandersee and Elisabeth Schussler, is at play. When people fail to see the plants around them, they do not notice their beauty. Worse, they undervalue their essential role in supporting all life on Earth. I wonder if some have become phytophobic, in their carelessness. As the world shifts from rural to urban and suburban living, we've become detached from plants, the very things that feed and oxygenate us. Even while many of us grow them, I fear we've also become alienated. To many, plants are just commodities, means to ends. I see it in the nursery catalogs we browse, the landscapes we design, the gardens we steward. We rush to buy the latest disease-resistant cultivar of elm, the hottest-off-the-press hydrangea, the newest, dazzling coneflower\u2014each a must-have due to superb pest-resistance, 5-season interest, drought tolerance, pollinator-friendliness, non-invasiveness, maximum winterhardiness, and a host of other traits folks claim the plants possess. We demand a lot from plants in our managed landscapes, and those qualities are essential for many to survive. Yet among the attributes of new garden plants, I see a glaring omission: a richly authentic connection to a human, something that makes each individual stem stand out among its green brethren. At the Arboretum, we do not grow trees: we jointly cultivate them and curate their narratives. Some of these are backstories that came with the plant; others emerge where they are growing. I may lecture to a class about Ernest Wilson's 1901 collection from China of the nowthreatened paperbark maple, Acer griseum; a docent may share with a visitor Connie Derderian's heroic efforts saving the bonsai and penjing collection while its curator from 1969 to 1984. This technique\u2014sharing their backstories\u2014 may be a surefire way to inspire others to see plants, but it still doesn't go far enough. The student returns home from the lecture hall; the visitor departs the Arboretum. Motivated as they may have been at the time, the moment can be fleeting. People need their own personal, authentic connections to plants, too. I'm lucky to have a garden at home, where I grow plants replete with backstories important to me. Don't get me wrong\u2014I grow quite a few eye-candy plants, too. Passersby from the sidewalk regularly ask the name of the shrub humming with bumblebees and awash with huge yellow blooms each July (Hypericum frondosum 'Sunburst'), and then follow-up wondering what that nearby mass of lacy, chartreuse vegetation is (Rhus typhina 'Bailtiger'). While they may be clickbait, these are not the real showstoppers. I get access to surplus seedlings from my own expeditions, and a dozen have found space at home. When I watch each bloom of Weigela decora shift from cream to rose, I recall that slightly rainy September day in 2018, collecting it with colleagues in Nikko, Japan. As I cut back the Hydrangea arborescens every spring, I harken back to 2014, gathering seeds in the Ozarks, shouting warnings to fellow collectors about the cottonmouths and rattlesnakes underfoot. I also grow a Stewartia rostrata collected by Peter Del Tredici in 2004, in Lushan, China. Every time I see it, I think of Peter and our friendship. None of these plants is the most glamorous of its kind, but I'd not trade them for the showiest of cultivars sold in fancy containers. I also grow plants passed down from family. A favorite is Rudbeckia laciniata 'Hortensia', often called golden glow, outhouse plant\u2014or its most colorful moniker, shithouse daisy. We called them \\\"grandpa's flowers,\\\" a name my mother grew up with, for her own grandfather dug them and other plants from the small farm in Hamlet, Indiana, loaded them in a horse-drawn wagon full of furniture, and drove the family to Sturgis, Michigan\u2014a three-day trip. Sure, the plant can be ratty, the leaves covered with mildew by the end of summer. But, when I see those bright yellow pompon blooms, I smile and think of generations of gardeners sharing the plant and the story. Plant lovers love to share plants. A decade ago, friends gave me fresh cuttings of a citronella-scented geranium (Pelargonium graveolens), which they in turn acquired as cuttings from a spontaneous plant sprawling on Mount Lycabettus, in Athens, Greece. My cuttings rooted and became houseplants that thrive under neglect. Last summer, I introduced the magic of plant propagation to one of the neighbor kids. Six-year-old Abe snipped a few stem cuttings, prepared and stuck them in a pot of basic soil (this geranium will root in anything!), and waited. They rooted, and a year later he has a lanky, smelly houseplant. Whether he tells the story of its Greek provenance matters little to me\u2014what does matter is that he has his own authentic backstory connecting him to the plant. By the end of his song, Kermit realizes that green is beautiful and exactly what he wants to be. Even when lacking fancy tradenames or cultivar epithets, the green beings in our gardens are beautiful when they have authentic backstories. As we grow the triedand- true (as well as novel-and-new) plants for ourselves, our clients, our cities, we must also make room for those with personal stories to learn, preserve, and share. If none come to mind, we write them by sowing a seed, rooting a cutting, or adopting a nearby street tree to call our own. In doing so, we will all better see the plants around us. They need this, and so do we."},{"has_event_date":0,"type":"arnoldia","title":"To Wander About","article_sequence":1,"start_page":1,"end_page":2,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25792","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15ebb6b.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Friedman, Ned","article_content":"To wander about among a vegetation which is new to one is pleasant and instructive. It is the same with familiar objects: in the end we cease to think about them at all. What is seeing without thinking?\nJohann Wolfgang von Goethe\nWe live in an age of ecosystems and genomes, where the scale of biology is usually presented at one of two extremes, global or genomic. There are good reasons for humanity's focus on the global scale of biology here and now in the Anthropocene. With human-induced climate change in the process of permanently altering the natural trajectories of nearly four billion years of evolution and ecological interactions between species, there is an intense focus on documenting and predicting what our single species has unleashed on the many millions of species of life with whom we have inherited and currently share the planet.\nAt the same time, the miracles of DNA sequencing technologies have allowed us to understand, in ways previously unimaginable, our own extraordinary evolutionary journey of becoming human, connecting us back in time to the first single celled forms of life. Reading the DNA has also provided amazing insights into everything from the genes responsible for making a flower to the genetic coding that maps out neural networks in fruit flies.\nIf one views the living natural world predominantly through the lenses of ecosystems and genomes, however, then something has been lost. I am an organismic biologist, a plant morphologist to be more precise. Simply put, this means that when I think of a 'unit' of biology, I am thinking about single organisms, just as you and I, as members of the human species, are single organisms. We are conceived as a zygote, develop into an embryo, are born, grow, learn to walk and speak, have interactions with other human organisms, and eventually complete an arc of life that returns our carbon to the earth. Of course, there is no such thing as a single organism all organisms depend on a web of myriad other species but I identify as an organism, knowing full well that there are roughly as many bacterial cells in my body as there are human cells. And the tree outside of my window, even though I know it has complex associations with mycorrhizal fungi and bacteria, is still a unit of biology that can powerfully be seen as an organism.\nI yearn to see organisms individual trees to meet them, witness them, learn from them, and indeed, to age with them. And this is the beautiful thing about the Arnold Arboretum and its roughly 16,000 accessioned woody plants. Each has provenance an organismic history with an origin story, and all that goes with siting, planting, and caring for an individual plant over decades and even centuries. I can reflect on the magnificent twisted European beech (14599*A) that was collected in the wild in France, transported to Royal Botanic Gardens, Kew, and then sent on a journey to the Arnold Arboretum in 1888. I can imagine a mere sapling being planted in the ground on the south flank of Bussey Hill in the beech collection. My mind reels in the magnitudes of time as I reflect on the generations of horticulturists who have cared for this one individual. And here, more than a century later, I can rejoice in its magnificent fall colors, its snow-covered spiraling branches, the light-green and delicate newly-flushed leaves in the spring, and the deep greens of summer. At the Arnold Arboretum, everything is truly about paying it forward.\nNot long after settling into the Arboretum, I resolved that I would never let a week go by without getting out onto the grounds to look at and photograph the woody plants that had beckoned me here. On every walk, I bring my small pocket camera and take pictures. Each night, I select the better ones, and spend additional time reflecting on what was revealed to me. By simply taking the time to observe, I feel as though I have gotten to know these non-sentient organisms on their terms: not as extensions of me, but rather as fellow living beings that can reveal their lives, history, complexity, beauty, architecture, and basic natural history.\nOver the years, from these meandering walks, I appear to have developed several of what I now refer to as (healthy) obsessions with phenomena which, once I observed them in the Arboretum, I became acutely interested in seeing in all of their manifestations. These obsessions include my ongoing annual spring quest to witness the brilliant hues of ovulate (seed-bearing) conifer cones; the exuberance of budbreak among the horsechestnuts and buckeyes; a fixation on the magical dispersal of pollen from rhododendron flowers; magnolias in fruit (and always, the bigleaf magnolia in flower); smooth bark (especially among snakebark maples in the winter); the startlingly bizarre naked resting buds of India quassiawood and the Arboretum's single specimen of Caucasian wingnut; looking straight up the trunks of large trees in all seasons; acorns in August (and the mad dash to finish filling up the fruit in the early fall); and the act of shattering birch infructescences to gaze upon their minute, delicately winged seeds, which immediately lift from my palm and are carried off by the wind.\nDiscovering plants as individuals, organisms to be reckoned with and reflected upon, is a journey worth taking, and one that never ends. It is a journey that enriches my life every day, in ways that I could not have imagined as I made my first focused perambulation on the grounds of the Arnold Arboretum years ago. It is also a journey that will be unique (and uniquely rewarding) to each person who sets out to discover the essence of plants by meeting these magnificent organisms on their terms, simply by looking and reflecting.\nIf we are ever to save the planet from our destructive tendencies, of warring with nature and each other, I would like to suggest that it can start by regularly walking in a garden, a park, the woods, or one's backyard, and learning to rejoice in the extraordinary beauty of organisms that can't talk to us, and indeed are wholly indifferent to our very existence (although certainly not unaffected) but whose presence is a constant reminder of the nearly miraculous complexity and interconnectedeness of life."},{"has_event_date":0,"type":"arnoldia","title":"Hedging Our Bets","article_sequence":2,"start_page":8,"end_page":10,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25791","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15ebb27.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Meyer, Mary H.; Kreevich, Nick","article_content":"Believe it or not, a hedge collection can be full of surprises. Take the row of 20-25 foot-tall Jack pines, Pinus banksiana, just one of the 73 taxa in the Hedge Collection at the Minnesota Landscape Arboretum (MLA). These tough natives have great cold and drought tolerance but what are they doing, unpruned, in this formal hedge collection? The historical documentation of the collection provided crucial clues: From early curatorial records, we learned the Jack pine hedge was made up of plants grown from seed collected by Al Johnson, an early MLA curator, from a witch's broom in Chittamo, Wisconsin. In slides, we found pictures of plant people with witch's brooms, from which they coveted seed in hopes of finding new dwarf plants. While the story of the Jack pines is clearer now, broad questions remain: What did record-keeping look like in 1967? Whose idea was it to start the collection? Was there an initial donor or collection goal?\nThese were some of our questions when we began to write our new ebook, Hedges: A Brief History and the Minnesota Hedges Collection (pressbooks.umn.edu\/hedges), and document the institutional knowledge of the collection. Although each plant had its own record with basic provenance information, our questions were not easy to answer. When we tell people (even horticulturists) that we wrote a book about hedges, they often look puzzled; when we ask if they grew up with a hedge, or if they ever pruned a hedge, however, most often the answer is yes. We ask them to think about why the hedge was there, and what it might have meant to their family and neighbors. And for many people, a light comes on as they connect their hedge to the landscape and its cultural meaning.\nEstablished in 1958, the MLA began as a horticultural research site for testing winter hardiness of plants (USDA hardiness zone 4), including plants commonly used in hedges. The MLA Hedge Collection is one of the oldest such assemblages, due in part to the boom in popularity that hedges saw throughout the 1960s, thanks to the postwar building boom and the growth of suburbia.\nSifting through thousands of more-than-60-year-old, 3 5-inch accession cards, filed in Steelmaster card cabinets, is like discovering an old journal or generational photo album at a yard sale: as soon as you start moving your fingers across the edges of the cards, musty whiffs of past time bring on a feeling of nostalgia. With each flip of a card, organized alphabetically by generic, specific, and cultivar epithets, you begin to build a historical portrait of the MLA collections dating from 1958. These index cards, also known as accession records, reveal that acquiring a plant and giving it a number did not necessarily coincide with when it was planted. Documentation on the cards also includes notes on fall color, winter injuries, fruit set, and overall growth habit all important considerations when assessing the ornamental value of a hedge. It is clear from the records that prior keepers prized foliage density, foliage color, and winter hardiness above all. We could even trace the impacts of weather on the hedge collection, with two of the coldest winters in recorded history (1978-9, with an average temperature of 9.4\u00a1F, and '77-8, with an average temperature of 10.5\u00a1F) apparent in plant-record notes on injury and severe dieback. Natural selection certainly took its course with those back-to-back weather events, but also provided the staff with critical knowledge of how particular hedge plants respond to extreme cold.\nWe also interviewed a number of employees, current and retired, to record their memories and discover the origins of some of the more unusual plants in the collection. Kathy Allen, Andersen Horticultural Librarian, assisted with locating the early Arboretum annual reports, which add critical details regarding scope of and support for the collection. 'The collection was planted to show which plants were the best for formal hedges,' recounts Director, Peter Moe, himself a longtime MLA employee. 'There were fewer compact forms of many species at that time and many people tended to try to keep large plants such as Amur maple as medium-sized hedges.' Height, density, and diversity could be shown in a planting a variety of hedges, which at the time were an extremely common and desirable landscape element.\nThree taxa of boxwood (Buxus 'Glencoe' Chicagoland Green\u00aa; B. microphylla var. koreana, and B. sempervirens) are the only broadleaved evergreens in the collection. Notes from early Arboretum newsletters express interest in this genus, though it was thought not to be winter hardy by many. Accession records and notes from the '60s to '70s document the overturning of this wisdom, with comments such as 'best in collection' and 'very good hedge material.' Although the plants show winter burn in the spring many years, the hardiness of boxwood hedges is no longer a question.\nGuided by prior documentation standards at the MLA, evaluating our current hedge collection for ornamental value sometimes felt like being a judge for the Westminster Kennel Club Dog Show, looking at one favorite after another. Usually, the first concern when pondering a hedge is its performance as a barrier. Yet not all hedges intend to create a barrier, but may instead provide ornamental value to an existing landscape or garden. The latter is mostly a matter of opinion: some may be wowed by the texture or height of a hedge, while others are more interested in the seasonal changes such as flower, fruit, and fall color.\nOur Thuja occidentalis 'Wareana' (American arborvitae) was the clear winner in terms of privacy, reaching heights of up to ten feet at maturity, with very dense, evergreen foliage. On the opposite end of the density spectrum, a hedge like Hydrangea arborescens 'Annabelle' (smooth Hydrangea) needs consistent renewal pruning to provide what it is best known for: its large, rounded flower heads. While the 'Annabelle' hedge can reach heights that garner it semi-private status, there is no hiding behind this deciduous shrub come winter. Deciduous hedges vary greatly as the seasons change, and will react strongly to weather anomalies. Early in the growing season, Philadelphus coronarius 'Aureus' looks like a candidate for removal, but its charm and value for use in a hedge come to fruition in the summer, with its beautiful chartreuse foliage.\nSuch factors are boldly visible through the seasons at the MLA, when you drive over the crest of a small hill and 73 neat hedges suddenly spread before you. It is hard to not notice them! Our hedge collection is a document of human intervention in the landscape. While it is doubtful that we would plant a hedge collection in a public garden today, there is value in keeping historical horticultural garden elements intact for future study. This preservation effort encompasses both plants and the records we keep of them.\nAnd so a word of advice to anyone thinking of fleshing out historical documentation for a plant collection: much institutional knowledge exists only in the memory of long term staff. Documenting this tacit knowledge, with audio or video recordings if possible, as well as continuing to keep records beyond mere accession numbers, will help curators, horticulturists, and other Arboretum staff understand the goal and educational purpose of a collection. Rarely do we over-document the details of our plant collections. As authors on hedges, our perspective has become more complex, realizing that people subconsciously use hedges to take control of their property and show authority. Some may balk at using the word 'authority' in connection with a hedge. Our reaction to plants is often subconscious, however; we rarely realize how deeply they affect us. A well-pruned hedge subconsciously communicates human control, and implies a safe, managed landscape. Our species' role in the landscape is readily seen, but often not fully recognized, when we encounter a hedge."},{"has_event_date":0,"type":"arnoldia","title":"Quest for Southern Red Oak-North of the Mason-Dixon Line","article_sequence":3,"start_page":11,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25790","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15eb76f.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Zale, Peter J.; Aiello, Anthony S.","article_content":"While plant collecting is often romanticized as occurring in pristine natural habitats, much of the most successful seed collecting is done in more prosaic locations. This is especially true when searching for tree species, where roadsides or power line rights-of-way provide light (for tree growth) and ease of access to fruiting branches (for collecting convenience). This certainly was the case in September 2020, as we searched for Quercus falcata (Spanish oak, or southern red oak) in southern Chester County (PA). After a fruitless morning at the Goat Hill Serpentine Barrens Preserve, we found much better success along local county roads.\nWhy Quercus falcata? Based on biological and climatic threats to tree species traditionally grown at Longwood Gardens, we have focused recent efforts on native species that combine ornamental traits with disease resistance and greater heat tolerance. For example, red oak (Quercus rubra), one of the more prominent shade trees at Longwood Gardens, in recent years has been among the most susceptible to bacterial leaf scorch (Xylella fastidiosa), which can weaken and eventually kill mature trees. As possible substitutes for this and other susceptible oaks, we targeted populations of Quercus phellos, Quercus michauxii, and Quercus falcata native to southeastern Pennsylvania. Maturing at approximately the same size as red oak, all three are potential substitutes as high-canopy, overstory shade trees. Each of these three species, which are widespread further south, reach the northern limits of their native ranges in southeastern Pennsylvania, southern New Jersey, and Long Island. Southern red oak barely extends its range into southeastern Pennsylvania, southern New Jersey, and possibly Staten Island and Long Island. In fact, due to its rarity in Pennsylvania (fewer than fifteen populations are known in the state), the Pennsylvania Natural Heritage Program lists it as a species of special concern (an S1, for those familiar with the coding).\nFaced with travel restrictions due to the COVID-19 pandemic, starting in the late summer and fall of 2020, we decided to look for local opportunities for collecting seed, pursuing a range of regional collecting objectives that we had previously not achieved. This included targeting the southern ranges of northern species (such as Larix laricina) and, vice-versa, the northern limits of southern species (for example Quercus virginiana), with the goal of growing plants suitable for changing climatic conditions in the Delaware Valley. For those southern species that reach into the mid-Atlantic, the extreme populations might possess traits, including cold tolerance, that provide opportunities to grow these beyond their traditional horticultural ranges. Conversely, for the northern species, the southernmost populations could possess greater heat tolerance, allowing us to continue to grow these as their native populations retreat northwards in the face of warming climates.\nIn September 2020, we focused our collecting on nearby populations of Quercus falcata, which occur on serpentine soils and their associated barrens. (Serpentine barrens, with thin, nutrient-poor soils, support high levels of unusual, rare, or endangered species, in contrast to adjacent areas.) We had targeted Goat Hill based on recent herbarium records from this location, but we did not find any southern red oaks there. Collecting was easy along the county roads, however, where we made three separate collections from trees whose acorns were within reach of our pole pruners. These three populations, within two miles of each other, were made up of large mature trees that were at least 50 feet tall. Quercus falcata stand out among other oaks, having coarsely lobed leaves with sickle-shaped (falcate) terminal lobes, and dense grey down (pubescence) beneath. For two of these collections, the southern red oaks were mixed among other native trees species in a dense forest; the third location was a grove of separate mature trees growing in a heavily grazed cow pasture, all within sight of the Herr's Snack Factory, a local landmark.\nSeeking additional Pennsylvanian locations of Quercus falcata, we pored through herbarium collections shared through the Mid-Atlantic Herbaria Consortium. Historical records from the late 1800s through the mid-1900s show a distribution in southeast Pennsylvania along much of the Piedmont-Coastal Plain boundaries, including southwest Philadelphia. But due to urbanization of much of this historic range (which includes Philadelphia International Airport), herbarium collections since the late 1980s center on three areas: southern Chester County near the Maryland border, southern Montgomery County along Militia Hill in Fort Washington State Park, and southern Bucks County along the Delaware River.\nWhile collecting southern red oak within a few square miles of West Nottingham Township (Chester County), we came across a remarkable diversity of eight oak species in addition to Quercus falcata, we also encountered Q. alba, Q. ilicifolia, Q. marilandica, Q. prinoides, Q. rubra, Q. stellata, and Q. velutina. Though familiar with the local diversity of oaks in southeast Pennsylvania, we rarely see this number of species in a single day's outing. Together, these represent a significant portion of the 11 oaks reported by Hugh Stone in his two-volume 1945 Flora of Chester County Pennsylvania, and nearly half of the approximately 20 oak species found in Pennsylvania. This wealth of oak species serves testament to the richness of the Chester County flora, historically the most botanically diverse in the state, though heavily impacted by human activities since the publication of Stone's Flora.\nWe returned in 2021 to make duplicate collections from the easily accessible roadside trees found in 2020. Oaks are famous for having years of heavy acorn production (mast years) followed by years of lower production. In 2020, we experienced a post-mast year when looking for Quercus phellos in southern Bucks County. Despite seeing a few dozen trees during a day in the field, we did not see a single acorn on any of these. Our luck was better with Quercus falcata: in 2020 we collected a total of approximately 250 acorns, and in 2021, just under 200 acorns.\nAs with seed collecting, patience is the main ingredient needed to grow oak seedlings. To germinate, acorns usually need a few months of cold treatment, followed by warm conditions and the increasing day lengths of spring. Ultimately the seedlings derived from these collections will be evaluated in our Research Nursery for growth rate and form, disease resistance, and fall color, before being introduced into public areas of Longwood Gardens."},{"has_event_date":0,"type":"arnoldia","title":"Hybrids Hiding in Plain Sight","article_sequence":4,"start_page":14,"end_page":15,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25789","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15eb36c.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Connolly, Bryan","article_content":"Last summer, while working as a consulting botanist for the Environmental Protection Agency's National Wetland Condition Assessment project in Allamuchy Township, New Jersey, I found an unusual colony of shrubby dogwood in the genus Swida (previously known as Cornus). The research plot was in a seasonally flooded meadow, with broad-leaved cattail (Typha latifolia), ironweed (Vernonia noveboracensis), and reed canary grass (Phalaris arundinacea) present. The site was previously cleared and looked to have a long history of human disturbance. The location is now set aside as town open space, and secondary succession is occurring, with woody plants increasing in dominance. Swida, or shrubby dogwoods, are known as old-field colonizers, and also as wetland species. There were two species present at this locality, S. racemosa, gray dogwood, and S. amomum, silky dogwood. In my experience, S. racemosa is more of an old-field colonizer, while S. amomum tends to favor wetlands. The area was both an old field and a wetland, and so it made sense that the species were co-occurring and abundant at the site.\nI noticed one Swida colony that did not cleanly fit into either S. racemosa or S. amomum. Swida racemosa generally has narrow (lanceolate) leaves, white fruit, gray bark, white pith, and upright growth habit; while S. amomum has broader (ovate) leaves, reddish-purple bark, brown twig pith, blue fruit, and a mounded growth form. The atypical plant I spotted had S. racemose characters, including narrow leaves 2.5\u00d03.8 cm wide and verrucose gray bark on the older stems, but also displayed the S. amomum traits of blue fruit and brown twig pith. Additionally, the growth form was unusual: it was a tall plant, about 2\u00d03 m in height, and somehow both upright and mounded, intermediate between the habits of S. racemosa and S. amomum. The pedicels or flower stalks were also reddish-maroon, not the typical bright red of S. racemosa. With this combination of characters, I thought it was likely to be a hybrid of the two species. From my experience working with coauthors on The Vascular Plants of Massachusetts: A County Checklist, I remembered a Swida hybrid, though I couldn't recall the parental species or if it was named. Additionally, from my wanderings and botanical work in the Northeastern US, I have published many new records of hybrid taxa and I could not place this plant among them.\nAfter a long day in the field I returned to my hotel room, fired up Go Botany (the online database of the Native Plant Trust), and confirmed that S. racemosa and S. amomum do in fact hybridize. On the account of my vague Swida hybrid recollection and my previous encounters with hybrid taxa, I wasn't surprised that a cross was known, but was glad that my hybrid hypothesis was supported by the literature. To my delight, the hybrid was listed as a nothospecies (a direct hybrid of two species) with the name Swida arnoldiana. The original description, by Alfred Rehder, was made in 1905 from a row of shrubs growing at the Arnold Arboretum.\nThis individual could just represent variation found within S. racemosa, which occasionally can have brown pith or light blue fruit. But I find it unlikely that a plant would exhibit both these traits while also co-occuring with plants that have the morphology of S. racemosa and S. amomum. I thus believe this plant to be S. arnoldiana. If I am correct, then it is a state record for New Jersey! According to Flora of North America, the hybrid has only been found in Massachusetts, Missouri, Ohio, and Pennsylvania.\nThe specimen voucher will be deposited at the Arnold Arboretum herbarium. This unique hybrid individual spotted in the field offered a nice little puzzle to solve and it was gratifying to learn that it is named after a wonderful arboretum I know and love!"},{"has_event_date":0,"type":"arnoldia","title":"Somewhere in the Panhandle of Florida","article_sequence":5,"start_page":16,"end_page":17,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25788","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15eb328.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Thomas, Elizabeth; Boland, Tim","article_content":"Somewhere in the panhandle of Florida, traveling for miles within a labyrinth of perfectly paved yet utterly empty roads, we blindly followed our guide, Bob, to a population of our target species, cracking jokes as we drove about the sinister fate awaiting us. We were in the ghost of a ghost town: the skeleton of a subdivision that was never built, planned for a population that never came. Every street is identical, save for the occasional cul-de-sac jutting into scrubby second-growth forest. Not only did the people never arrive, but the houses were never constructed; hundreds of miles of paved roads were laid here in the 1990s, only to be left abandoned, another suburban dystopia created by Florida's rich history of speculative development.\nBob pulled his Prius to the side of the road, and we parked behind him. We'd been put in touch with Bob, a local resident and active amateur botanist, by our contact in the Florida Forest Service; it had been Bob who discovered this population of Stewartia malacodendron, and he was eager to share it with trustworthy enthusiasts. Known commonly as 'silky stewartia' or 'silky camellia' due to its showy, camellia-like blooms, S. malacodendron is a small deciduous understory tree native to the southeastern coastal plain of the United States, from Virginia to the northeast and Texas to the southwest. Traveling from the Polly Hill Arboretum on Martha's Vineyard in the late summer of 2021, we had come to harvest fruits (and thus, seed) of this species to grow into plants for our ex-situ living collections. As co-holders of the Plant Collection Network's National Collection of the genus Stewartia with the Arnold Arboretum, we were hoping to collect from populations at the southernmost edge of its range, heretofore unrepresented in our living collections.\nGrabbing our gear, we ambled through the woody goldenrod and beautyberry, enjoying the fresh light of an early September morning pouring through the sparse canopy of southern magnolia and live oak. We were only about 100 feet from the road when we stumbled across our first stewartia tree, standing like a sentinel at the rim of a large, sunny slope dropping down to a sinkhole pond. Our satisfaction to find it fruiting quickly turned to excitement and then overwhelm as we spotted at least a dozen more of them spread across the face of the slope, each one dripping with plump, green fruits larger than we had ever seen before. Normally about an inch in diameter, these were more akin to fuzzy ping pong balls or small crabapples. Balancing on the scrubby slope and madly scribbling collection numbers on sandwich bags, we exchanged involuntary expletives as we took in the superlative bounty of fruits. Were we harvesting germplasm, or were we apple picking?\nOf conservation concern throughout most of its range, silky camellia is a protected endangered species in the state of Florida. Protected in theory, at least: as we began to stuff the plump green fruits into plastic bags, we gazed across the sinkhole, where a collage of zig-zagging tire tracks defaced the far slope all the way down to the shore of the pond. With its bleached, eroded sands, this local party spot is visible even from satellite images.\nLiz heard Bob holler from somewhere upslope to her left. Though she couldn't make out what he was saying, she knew he'd found yet another group of exceptionally fecund trees. This good news found her in a fluster of multitasking, as she scrambled to capture location coordinates on our GPS unit, measure and record specimen data and collection numbers by hand in our field book, label baggies and herbarium samples, take pictures, and collect fruit. Sharpie cap in mouth and hair sticking to her sweaty forehead, she wondered whether we'd be late to our next stop that morning, meeting our next guide at a site about three hours west. We'd expected this to be a quick roadside stop, not an absolute windfall.\nThis expedition is our most focused effort to collect this species since an Alabama excursion in 2007. Our founder, Polly Hill, was among the first private collectors to grow this plant, with our oldest tree dating back to 1962. The mild maritime climate and acidic soils of Martha's Vineyard happily support the cultivation of this stunning, small, flowering tree.\nMost of our expedition planning is done months ahead to arrange for a seasoned naturalist or professional botanist to lead us to target species. Admittedly, we get into some very wild places to collect the silky stewartia. Usually, we find them after rugged hikes into deep wilderness. Yet here on this fine morning, just off an intersection crossroad, we had found the most robust population of silky stewartia Tim has witnessed in 15 years of pursuing it. Slicing into the globose fruits, we found dark brown sclerified seeds that shone brightly in the late morning light.\nThis unexpected and surreal discovery was a vivid reminder that plants are both resilient and vulnerable. The silky stewartia is imperiled by habitat destruction, principally through logging or building development. On a previous scouting trip to Alabama in 2012, we bore witness to a new condominium development that destroyed a former thriving population. Somehow, this spectacular hillside of trees was spared the backhoe and bulldozer for now. With the same luck that brought us to this unique population of trees, we hope to return and see them in bloom someday. Perhaps the flower size will also be larger, or some of the petals streaked ruby red, as variants in the wild are known to do?\nWe looked at our watches to check our time; not surprisingly, we would be late to our next destination. However, the place, the trees, and the experience were worth it. As we gathered up our gear to move onto our next location, we did so in a suspended state of stewartia euphoria. The remainder of the trip was both productive and satisfying, but nothing would compare to this remarkable discovery in the unlikeliest of places."},{"has_event_date":0,"type":"arnoldia","title":"Rhododendron prunifolium","article_sequence":6,"start_page":18,"end_page":19,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25787","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15eaf25.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"I have always been fond of Korean azalea (Rhododendron mucronulatum), that hardy shrub whose pink flowers crack open just as the snow recedes. At the other end of the season, and the last of our azaleas to flower, comes another personal favorite: Rhododendron prunifolium (plumleaf azalea).\nMost of the Arboretum's plumleaf azaleas grow along Meadow Road, amassed below towering black locusts in the Wolcott Bed. They escape notice until the middle of July, when their floral buds swell and burst open, when few woody plants bloom and temperatures are irrepressibly hot. Although it's not comprehensively accurate (color ranges from deep red to nearly pinkish-orange), in this portrait, I'll say the flowers are safety orange, that alarming shade reserved for prison jumpsuits and cautionary traffic cones.\nThis color should be taken as a warning, because Rhododendron prunifolium is rare in nature, limited to just a few dozen populations in the Chattahoochee River Valley and straddling the border of southern Alabama and Georgia. Neither disease nor insect is to blame; climate change (to date at least) is also not the culprit. Instead, the species totters on the brink simply because its preferred habitat mesic forests, stream sides, and ravines is disappearing due to logging and other development. In this respect, plumleaf azalea is like most other woody plants threatened with extinction: their natural homes are vanishing.\nShortly after the founding of the Center for Plant Conservation in 1984 (then based at the Arnold Arboretum), we began to collect the species in earnest. At present we grow thirty-four plants, mostly from Georgia's Dade, Harris, and Stewart Counties. Preserving wild populations remains the highest priority, but it is important to have a back-up; while they grow here, their story is shared with others, and scholars from around the world come here to study them.\nLet's not ignore the fact that Rhododendron prunifolium looks good in the garden. For endangered species, being charismatic and attracting attention is a gateway to its security (just look at the giant panda). This means we must equally care for species whose security is questionable simply because they are less charming, at least in appearance anyway. At the Arnold, we make room for these plants, too."},{"has_event_date":0,"type":"arnoldia","title":"Beatrix Farrand on Mount Desert Island","article_sequence":7,"start_page":20,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25786","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15eab6d.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Eason, Rodney","article_content":"I first visited the Abby Aldrich Rockefeller Garden, in Seal Harbor, Maine, on vacation with my then-fianc\u017de, now-wife, Carrie, in 1997. We were both young landscape architects practicing at different firms in Raleigh, North Carolina. The garden visit had been arranged by Carrie's college classmate, Sarah Richardson, who lived on Mount Desert Island. After days spent hiking through Acadia National Park's coniferous forests, granite peaks, and scattered blueberries and junipers, the refined curation of color within the borders of the Abby Aldrich Rockefeller Garden was a beautiful and dramatic contrast.\nSarah informed us that the Rockefeller Garden was designed by Beatrix Farrand (June 19, 1872-February 28, 1959), who also had designed Dumbarton Oaks in Washington, DC. One of Carrie's classes at Penn State had made a trip to our nation's capital, where she had been awestruck by the beauty of that garden. The only images I had seen of Dumbarton Oaks came from books and slide lectures, and it would be roughly 18 years before I would encounter Farrand's work in depth, reading her biography by Judith Tankard, Beatrix Farrand : Private Gardens, Public Landscapes (2009).\nToday, through a set of fortunate circumstances, I get to live all year round on Mount Desert Island and have served as CEO since 2015 of the Mount Desert Land & Garden Preserve, which is entrusted with the care of three Farrand-influenced gardens, including the Abby Aldrich Rockefeller Garden. In case you have never visited the coast of Maine, I should point out that the indigenous vegetation is not exactly floriferous. Coniferous forest predominates, largely composed of red spruce, black spruce, and white pine. There are some deciduous trees on the edges of the coniferous stands, including alders and moosewood maples. The dominant native ground-floor vegetation is largely composed of rhodoras, sweet ferns, huckleberries, blueberries, and northern bayberries. Underneath this typical plant community on Mount Desert Island are numerous ferns, mosses, lichens, and sedges. This plant community makes for a mix of greys and greens, all in contrast to the pink granite outcrops and glacial erratics that you would frequently encounter. Spectacular in its own right, this landscape inspired the formation of Acadia National Park in 1916.\nWith English-style, mixed-herbaceous borders set off within this landscape, the Rockefeller Garden makes an evocative juxtaposition. Designed by Farrand for Abby Aldrich and John D. Rockefeller Jr. from 1926 through the early 1930s, the garden is a sublime mixture of sophisticated design and a complex palette of plants. I was smitten from the outset with the combination of bold floral colors, statuary sourced from Asia, and Beaux-Arts symmetry, provided most prominently by two parallel axes that run the length of the garden and orchestrate the flow for the visitor. The entry axis, called the Spirit Path, is flanked by carved-stone warriors and priests from eighteenth-century Korea. The second axis, parallel to the Spirit Path, provides the central aspect of the garden and its colorful flower borders with a distant view of a round opening called the 'moon gate.' This gate frames the view of a eighteenth-century bronze Buddha in the Shakyamuni, or historical form, from China. As I studied Farrand's designs in more detail, I would learn how the use of such central orienting axes became a hallmark of her designs.\nMany years after visiting Mount Desert Island and the Rockefeller Garden for the first time, I was fortunate enough to visit Dumbarton Oaks. Like an unfolding, complex novel that you just cannot put down, the garden kept leading from one masterfully designed room to the next, with brilliantly placed plants and sublimely scaled spaces. I distinctly remember encountering the camouflage-print bark of a superb Chinese quince, Pseudocydonia sinensis, at the end of a pathway. Overwhelmed by the beauty of this gorgeous tree, I walked off the pathway and gave it a hug.\nWhat is now the Land & Garden Preserve, was conceived in 1970 as a way for Peggy and David Rockefeller to perpetuate the beauty of the Abby Aldrich Rockefeller Garden. They co-inherited the garden with David's older brother, Nelson, after David's father, John D. Rockefeller Jr., passed away in 1960. Soon after Peggy and David formed the Preserve as a non-profit, then known as the 'Island Foundation,' they were asked to manage the nearby Asticou Azalea Garden. Asticou, or the Azalea Garden as it is known locally, had been built beginning in 1956 by Charles K. Savage, using mature plantings from Reef Point, Beatrix Farrand's Bar Harbor estate.\nAnother local garden, Thuya, joined the Preserve in 2000, after its trustees decided that the future of the garden would be in good hands with the growing organization. Thuya's origins date to 1912, when a Boston landscape architect and Northeast Harbor summer resident, Joseph Curtis, constructed his 'rusticator' lodge in Northeast Harbor, naming it for a prominent stand of eastern white cedar, Thuja occidentalis, growing nearby. Charles K. Savage became the trustee of Thuya after Curtis' death in 1929. In 1956, Savage began establishing gardens at both Thuya as well as Asticou, a story for which more detail will be provided below.\nTo celebrate his 100th birthday in 2015, David Rockefeller gifted the Preserve over 1,000 acres of land around Little Long Pond, including over 10 miles of carriage roads and 10 miles of hiking trails. This parcel, too, carried Farrand's legacy: When John D. Rockefeller Jr. was constructing the carriage road system from 1913 until 1940 on what is now both the Preserve and Acadia National Park, Farrand had provided pro-bono consulting on road layout and planting designs. When David Rockefeller passed away in 2017, the Farrand-designed Abby Aldrich Rockefeller Garden joined his gift to the Preserve. Beyond the beauty of the Rockefeller estate in Seal Harbor, Maine, she left an indelible mark within what is now Acadia National Park and the Preserve.\nHer ties to the place were deep. When Farrand was 10 years old, in 1882, her parents bought an ocean-front property called Reef Point in Bar Harbor, facing Frenchman Bay. Her childhood at Reef Point fostered a love of plants and landscapes, and for amusement she dug and transplanted native vegetation from the surrounding forests and combined these with cultivated ornamentals. Farrand's ethos of protecting the natural environment while cultivating intensive gardening spots of horticultural pleasure carries on today at the Preserve with over 1,200 acres of conserved, natural lands connecting our three ornamental gardens.\nAs her interest in landscape design and planting became more of a passion, she was introduced to Charles Sprague Sargent, the first director of the Arnold Arboretum. Sargent agreed to guide Farrand in her self-education in horticulture and garden design from 1893 to 1894, since at that time, no formal training in landscape architecture existed. While studying at the Arnold, she worked with plants at the Arboretum, as well as at the Sargent family's estate, Holm Lea, in Brookline, Massachusetts. In addition to learning about the art and science of horticulture from Sargent, she learned to design landscapes to fit a site rather than change a site to fit a design.\nThe lessons she learned from Sargent carried over as well through the trialing of new plants at Reef Point and elsewhere. From 1946 to 1956, Farrand chronicled the evolution of her Bar Harbor garden along with the noteworthy characteristics of many plants in the 'Reef Point Gardens Bulletin.' Farrand found the climate of Mount Desert Island to be particularly hospitable to climbing vines and in the June 1954 bulletin, she describes some of her favorites. Among her descriptions of Aristolochia spp., Ampelopsis brevipedunculata, Vitis spp., and Lonicera spp., Farrand is particularly hopeful and enamored by a vine that 'Professor Sargent had scornfully described as a 'dud.' This Arnold Arboretum cast-off was Tripterygium regelii. I admittedly had never heard of this Celastraceae member until this mention in the Reef Point Bulletin.\nWhat began as a joint venture with her husband, Max, Farrand continued to develop, seeking to make Reef Point a public teaching garden after his passing in 1945. Max had been the first director of the Huntington Library and Gardens in San Marino, California. The Farrands divided their time between San Marino and Bar Harbor, with a dream of eventually making Reef Point a garden where aspiring horticulturists and garden designers could learn. In October 1947, two years after Max's passing, a massive wildfire burned almost a third of Mount Desert Island, including many grand, oceanside estates. These massive estates had provided the town with substantial tax revenues, now lost to fire. I mention this because Farrand had sought tax exemption of Reef Point as a public garden, and after these fires (which left Reef Point unscathed), the town had to increase tax assessments. With the burden required to keep her gardens afloat, Ms. Farrand ultimately decided to dissolve Reef Point as a lasting garden in 1955.\nBeatrix Farrand's article on 'The Azalea Border' in the April 15, 1949 edition of Arnoldia described the addition of azaleas and other acid-loving plants along Meadow Road by the Arnold Arboretum. Some of the azaleas mentioned in the article included: Rhododendron mucronulatum, R. dauricum, R. canadense, R. vaseyi, R. schlippenbachii, R. arborescens, R. viscosum, R. nudiflorum, R. roseum, and R. calendulaceum. After reading this article from 1949, I began to wonder if Farrand's interest in azaleas was in any way linked between her desire to see what would grow both in Jamaica Plain as well as at her Bar Harbor estate. As I will describe later, many of her plants were subsequently moved from Reef Point to the Asticou Azalea Garden and Thuya Garden by Charles K. Savage. Asticou Azalea has a substantial collection of azaleas, many of them species grown in the Arnold's Azalea Border. Farrand, along with her plant recorder, Marion Ida Spaulding, kept an herbarium of the Reef Point plants. Once Farrand decided to no longer keep Reef Point Gardens going, she sent their plant vouchers to the University of California, Berkeley's herbarium, where I have found 51 vouchers attributed to Reef Point.\nIn 1956, Farrand sold Reef Point to a Maine colleague, Reef Point board member and architect, Robert Patterson, who sold most of the plant collection to Northeast Harbor hotelier and fellow Reef Point board member Charles K. Savage. Lacking the $5,000 needed to purchase and move the collection, Savage was able to convince John D. Rockefeller Jr. to become a financial backer (that $5,000 in 1956 would be worth over $51,000 in 2022). Rockefeller and his wife, Abby, had worked with Farrand for over a decade on the design and construction of their Maine garden, what is now known as the Abby Aldrich Rockefeller Garden. Documents in the Rockefeller family archives show that many of the drawings for the garden and planting designs were by Farrand. After a trip to China in 1921, Abby Rockefeller became enamored with the pink stucco wall around the Forbidden City in what is now Beijing. It served as the inspiration for the wall that surrounds the Abby Garden in Maine.\nOutside of her formal garden designs, Farrand often acted as a consultant to Rockefeller about aesthetic decisions regarding the carriage roads both during and after construction. In correspondence in the Rockefeller Archives Center in Pocantico, New York, Farrand commented that the engineers and tradesmen that Rockefeller had hired to landscape the carriage roads of Acadia National Park were lining trees up like soldiers. She urged Rockefeller toward a more natural arrangement, mixing conifers and deciduous trees of different species and sizes.\nFarrand understood that the natural character of the carriage roads through the park required a more relaxed style than was evident in her notable formal garden designs. In other writings and sketches to Rockefeller, Farrand suggested covering many of the stone bridges with vines such as Parthenocissus quinquefolia (Virginia creeper). In Acadia National Park today, you will find 16 stone bridges built by Rockefeller, none of them covered with vines. Last year I was hiking along Stanley Brook on the southeastern side of Acadia National Park, and I stopped to admire the Stanley Brook Bridge. I noticed that at both ends of the bridge, equally spaced, was a pair of sugar maples. Growing four sugar maples so symmetrically, on both sides of the bridge, would have been a profound work of art for Mother Nature, so I am going to put this down to Farrand at the very least, a reflection of her influence and love of symmetry.\nOnce Charles K. Savage, or 'C.K.' as he was known locally, was able to secure the $5,000 from John D. Rockefeller Jr. for moving the plants from Reef Point to Asticou Azalea and Thuya, he had to act quickly. Savage wrote a narrative to Rockefeller, describing the need for funding and urgency in the matter. The new owner of Reef Point, Robert Patterson, was now responsible for paying taxes on the property and wanted Savage to move the plants before the property would be sold again. The move was done quickly, and records for which and how many plants were relocated remain elusive. White & Franke Tree Service, of Brookline, Massachusetts, with the assistance of various local helpers including Savage's young daughter and son, moved as many plants as possible the 11 miles from Reef Point to Thuya in Northeast Harbor. The Preserve has several historical photos of these plant moves; we thus know that White & Franke assisted with the move, as their company name is on the door of the moving truck. These photographs show that the largest plants were hand-dug, balled and burlapped with drum-laced jute, and moved with what looked like a converted tow truck, the lift on the back of which acted like a small, mobile crane. The plants were healed in and surrounded by mulch at Thuya during the winter of 1956, while construction continued at Asticou with the hauling in of truckloads of soil and stones that would eventually form the framework for the garden. Construction continued at Asticou and plants were moved from their temporary locations at Thuya until the garden was completed in 1958. Savage had also selected plants from the Reef Point collection for Thuya, planted after the Asticou plantings were completed.\nI hoped there was a document to be uncovered in someone's basement, outlining all the plants purchased, moved, and planted by Savage. During research for this article I learned that even Farrand was unsure of what existed at Reef Point. As she was building the collections, she noted her continuous desire to correctly identify the plants in the garden, even bringing William Judd, the Arnold Arboretum's chief propagator, to Maine for help with inventorying the collection. Whether due to the rapid movement of the plants, the transfer of records and herbarium vouchers from Reef Point to Berkeley per Farrand's request, or the inadequate identification of the plants by their owner, the Preserve has never had a consolidated record of what was moved from Reef Point to Asticou and Thuya.\nThe current manager of Asticou Azalea Garden, Mary Roper, has worked to identify the plants under her care for over three decades, including some of the plants moved from Reef Point in 1956. Mary began working at Asticou in 1989, some thirty years after the moves were completed. Over the years, Mary, like Farrand before her, has assessed the nuanced details of flowers, leaves, and stems of the plants under her care to develop a proper identification. Beginning in late 2022, Grace Brown, the Preserve's plant recorder and lead gardener at Asticou, will begin sharing some of these plant records via our new plant records database, accessible at gardenpreserve.org.\nDespite the remaining mysteries, the spirit of what Beatrix Farrand envisioned at Reef Point lives on today at the Preserve, within the gardens of Asticou, Thuya, and Abby Aldrich Rockefeller, as well as in the forests and meadows of our natural lands. This is felt most powerfully at the Abby Garden, with its overall layout, plantings, and ornamentation preserved since the 1920s. Asticou and Thuya were designs of C.K. Savage, but it was the influence of Farrand's relocated plants that completed these garden arrangements. When I tell someone who has visited the Preserve that I work there, 'I just love (insert either Asticou Azalea, Thuya, or Rockefeller Garden here)!' is usually one of the first things I hear in response. When I ask why they love their garden of choice, the responses often embrace the spirit of these places. I felt that special spirit when I first visited the Abby Garden in 1997, and I still sense this every time I visit. When I walk through Thuya, as I brush up against the old Kalmia latifolia that came from Reef Point, a quiet, distinguished vibe seems to emanate from the plants that came from Farrand. Asticou Azalea's design and plant masses are calm and subdued, much like I assume Farrand was during her life. Yet during the spring when the azaleas and cherries burst forth with an explosion of blooms, I can see Farrand's love of beauty in plants and the art of arranging a garden for others to enjoy."},{"has_event_date":0,"type":"arnoldia","title":"The Transatlantic Arboretum in the Nineteenth Century","article_sequence":8,"start_page":32,"end_page":41,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25785","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15ea76a.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Elliott, Paul","article_content":"In the summer of 1850, Andrew Jackson Downing embarked on a trip to England, where he toured gardens and rural estates. Downing was then thirty-four years old and had already emerged as a leading American landscape designer and horticultural writer. On the trip, he made a special stop in the midland town of Derby to see a garden known as the Derby Arboretum. The eleven-acre arboretum had been established ten years before, on land given for that purpose by a wealthy local cotton manufacturer, Joseph Strutt. Each tree was clearly labelled, and the arboretum, for two days a week, was completely free and open.\n'As a public garden'the gift of a single individual' it is certainly a most noble bequest,' Downing wrote. 'I met numbers of young people strolling about and enjoying the promenade, plenty of nurses and children gathering health and strength in the fresh air, and, now and then, saw an amateur carefully reading the labels of the various trees and shrubs, and making notes in his memorandums-book.'\nThe Derby Arboretum was distinct for its commitment to the public'even providing access to books so that interested visitors could learn more about the plants growing in the landscape. This commitment, Downing was sure, meant that the Derby Arboretum 'is, and will be, one of the most useful and instructive public gardens in the world.'\nOften considered the first public arboretum, it was designed by the Scottish landscape gardener John Claudius Loudon, who was most responsible for popularizing the term and concept of 'arboretum' during the nineteenth century. Yet public tree collections like those at Derby, and the Arnold Arboretum in the decades to come, did not arise de novo. Rather, the development of these institutions in Britain and the United States during the 'long' nineteenth century (encompassing the period from 1780 to 1919) is best understood as a global' and particularly transatlantic'phenomenon, arising at a time of large-scale immigration, industrialization, and botanical exploration. In that sense, public arboreta were products of changing relationships with the environment and, indeed, among people.\nOrigins of Transatlantic Arboreta\nThe Atlantic world was fertile ground for the formation of tree collections in the parks and gardens of Europe and North America. The vast forests of North America, with their seemingly boundless numbers of trees (many new to European science), inspired the formation of tree collections in those places beginning in the eighteenth century. The biodiversity of the North American forests spanned from subtropical to boreal, from coastal to montane. This diversity across the vast extent of the continent persists to this day, as exemplified by the ninety-nine native species of conifers now believed to exist north of modern Mexico. By contrast, Britain and Ireland have only three native species of conifers'and, in general, far fewer native trees.\nTransatlantic arboreta arose from a combination of tree collecting for gardens and parks and systematic planting in physic (i.e. medicinal) and botanical gardens. American trees themselves played a large part in this process, and were often collected in places known as 'American Gardens' between around 1700 and 1840. The enthusiasm for collecting American trees was encouraged by publications such as Mark Catesby's Hortus Britanno-Americanus, published in 1763, which emphasized the value of these plants for timber, shade, fragrancy, and beauty, holding them superior to British trees. Many American trees and plants were brought over to Britain and Ireland in the colonial period and early decades of the United States, especially through the botanist and explorer John Bartram, who, in the mid-eighteenth century, sent many examples to the English botanist and gardener Peter Collinson. Settlers in the New World also brought numerous trees from'and via'Europe with them, bringing these and trees from eastern North America with them as they moved westwards towards the Pacific during the nineteenth century.\nPlant collectors like Bartram were crucial to the creation of transatlantic arboreta, and came to be seen as heroic figures, making expeditions on behalf of wealthy collectors, nursery companies, governments, and scientific institutions. In his Dendrologia Britannica, published in 1825, the Hull merchant and botanist Peter William Watson praised the 'bold and scientific travellers' traveling throughout North and South America and other regions and identifying thousands of species. One of the most famous plant hunters of the era was the Scotsman David Douglas, who trained at the Glasgow Botanic Garden and made three separate collecting expeditions to North America in the first half of the nineteenth century. His introductions into Britain from the West Coast included the Sitka spruce (Picea sitchensis), Douglas fir (Pseudotsuga menziesii), red alder (Alnus rubra), and many others.\nNursery companies in America and Britain came to specialize in obtaining and selling American plants. The Loddiges company in Hackney, London, for example, had an American Garden, and featured many American trees in their collections and sales catalogues. Loudon used their collections for his research. Wealthy British aristocratic collectors such as the Duke of Devonshire at Chatsworth'a landscape designed by Joseph Paxton, a noted English designer'spared no expense in obtaining 'exotic' trees from America and across the globe for their parks and arboreta with the same eagerness that they acquired antiquities and works of art.\nMeanwhile, in North America, a series of private gardeners began to establish systematic tree collections, although they were not always designated as arboreta. For instance, William Hamilton, a neighbor of the Bartrams, developed his estate known as the Woodlands on the Schuylkill River, then outside of Philadelphia. In the decades following the Revolutionary War, he formed what was then one the largest American tree collections, arranged in the style of an 'English garden.' He toured gardens in Europe and obtained specimens from the Chelsea Physic Garden in London and other international sources. Private collections like this would inspire public institutions to come.\nLiving and Paper Arboreta\nOther inspirations for Atlantic world arboreta were the publication of arboriculture books, which were, in effect, 'paper arboreta.' The writing was informed by living tree collections. General studies of arboriculture grew from classic tree studies such as John Evelyn's Sylva, or a Discourse of Forest Trees, from 1662. The popularity of works such as Erasmus Darwin's epic poem The Botanic Garden, published in 1791, demonstrated how systematic plant collections were gateways to enchanting and exciting scientific worlds. The poem was initially inspired by a botanic garden Darwin established near Lichfield, England, which successfully united landscape beauty with Linnaean botany'and the book was much reprinted in British and American editions.\nHorticultural periodicals such as Loudon's Gardener's Magazine and Downing's The Horticulturist (first issued in 1826 and 1846, respectively) helped build public enthusiasm for trees and landscapes. Both men advocated for the development of arboreta as part of suburban gardens. The collections could be associated with park development or collectively give the appearance of a country park through combination of private gardens, especially in the United States, where there were fewer walls and fences in between plots. Though space for such collections was sometimes limited, especially in Britain, Loudon argued that arboreta were ideal for middle class gardens, even for small houses and gardens.\nMoreover, a series of books on regional and national arboriculture provided lists of hardy British and North American trees and shrubs, contributing to the acquisition and collection of trees. The plants delineated in these publications often came from all over the world, and they were only 'British' or 'American' to the extent that they had proven hardy enough to be grown outside in those places. Watson's Dendrologia Britannica, for example, provided 103 plates of North American trees imported to Britain, alongside others from Southern Europe and West Asia.\nOne of the most influential of these paper arboreta was Loudon's eight-volume Arboretum et Fruticetum Britannicum, from 1838, which inspired the creation of many tree places, including the Derby Arboretum. It was, in many ways, a transatlantic work that drew on arboricultural literature and catalogues from across the Atlantic world to provide a detailed history of trees and shrubs from antiquity to the 1830s . According to William Jackson Hooker, director of the Royal Botanical Gardens at Kew, Loudon's study was a work of 'vast importance' not just to Britain and Europe, but also to 'the temperate parts of North America.' Loudon made full use of a transatlantic network of botanists, gardeners, nurserymen, landowners, and plant collectors who provided him with information and drawings, specimens, seeds, and other tree parts.\nThe first volume of the Arboretum Britannicum included a chapter on American arboriculture informed by American contacts such as the printer Colonel Robert Carr, in Philadelphia, who, with his wife Ann Bartram Carr, had taken over responsibility for maintaining Bartram's Garden. Loudon believed that although American trees and shrubs had been available in British nurseries for decades, many remained under-appreciated, and he hoped the Arboretum Britannicum and the living arboreta it inspired would increase the number and popularity of more public tree places showing off 'living specimens' and capturing imaginations in a way dried herbaria never could.\nPicturesque Naturalism, Tree Planting, and Arboreta\nTrees were also essential to transatlantic conceptions of landscape design, providing beauty, color, contrast, structure, variety, seasonal change, and much more. The dominant Atlantic-world landscape philosophy of the nineteenth century was known as 'English' picturesque naturalism. This style idealized the English landscape, and was widely invoked in garden, park, and arboretum designs. Downing, for example, believed that the style developed in Britain by the English landscape gardener Humphry Repton, Loudon, and others should be applied across North America. According to Downing's pupil and friend the Ohio landscape gardener Frank Jesup Scott, who published a popular book on suburban gardening in 1870, 'compared with the English' the Americans were still 'novices in the fine arts of gardening' and the 'exquisite rural taste' even shared by 'the poorer classes' of England.\nPicturesque naturalism encouraged the positioning of trees and shrubs to achieve effects of openness and simplicity, shelter, shade, and beauty, to obscure boundaries through screen plantings, and to offer the occasional pleasure and sublimity of distant views. The designs often emphasized varied sensory experiences: sloping and terraced ground, shifting light patterns, the sounds of leaves and water, and the changing colors and aromas of trees and floral displays. The movement of birds and wildlife added to this multivarious experience for visitors, especially to the extent that animals (like plants) had their own degree of controlled agency.\nFurther development of this transatlantic landscape gardening philosophy was encouraged by immigration and the movement of people across the Atlantic. British and Irish gardeners and landscape gardeners working in North America brought ideas and methods from home which they adapted to local conditions and contexts. Notably, while Downing was on his British tour in 1850, he met the architect Calvert Vaux and persuaded him to immigrate to America, joining Downing's practice in Newburgh, New York. In the decade to come, Vaux, a Londoner, would employ picturesque naturalism when planning of New York's Central Park, which he codesigned with Frederick Law Olmsted.\nThe careers of Vaux, Downing, and Olmsted, and their many other professional interconnections, illustrate how an international approach to designing with trees took root on both sides of the Atlantic. In the second half of the century, Olmsted became a leading practitioner of picturesque naturalism. Successful picturesque landscapes, according to Olmsted, worked by adapting and evoking nature to produce a 'higher impression of grace than nature minus the agency of man would have produced,' stimulating the 'simplest, purest and most primeval' actions of the poetical side of 'human nature,' offering relief from the overly elaborate but stressful 'sophisticated and artificial conditions of their ordinary civilised life.' In practice, of course, the features held to constitute this language or tradition underwent considerable variation, although it remained particularly important to many North American and British landscape gardeners to claim to be following this tradition. While there was some introduction of formalism and Italianate features from the 1850s and 1860s, the languages of picturesque naturalism remained highly influential throughout the century.19\nArboreta as Public Institutions\nThe appearance of nineteenth-century public parks and arboreta was associated with the development of modern urbanization across the Atlantic world with its new institutions, suburbs, transport systems, built environment, and cultural experiences. Travelers, books, and ideas crisscrossed the Atlantic, encouraged by more rapid and cheaper steam ship lines and technological improvements such as telegraphy and undersea cables. While immigration to North America brought immeasurable human resources, it also increased tensions, clashes of identity, and problems of health and sanitation in towns and cities. As the pattern of immigration changed, bringing new peoples from across the globe, the question of how to adapt British and European landscape gardening ideas and practices to American contexts became more contentious. However, public parks were promoted as rational recreational institutions which could help facilitate assimilation, intercourse between the classes, and American patriotism.\nIn the United States, some of the earliest tree collections in designed public landscapes were associated with suburban garden cemeteries or 'rural cemeteries.' Mount Auburn Cemetery in Cambridge, Massachusetts, was established in 1831 and soon followed by others, including Laurel Hill Cemetery, in Philadelphia. The cemeteries represented the application of landscape gardening aesthetics and practices. In London, Abney Park opened in 1840 and included collections that were formally laid out, at least in part, as a labelled arboretum. The landscapes were portrayed as sacred places where family members and others could repose in quiet contemplation amidst appropriately somber planting, particularly yews (Taxus), Scots pine (Pinus sylvestris), and other evergreens and columnar trees associated with mourning.\nEncouraged by Loudon in particular, a series of public and semi-public arboreta were established in Britain from the 1830s, while many new public parks and botanical gardens also featured arboreta. Arboreta were opened at Derby (1840), Nottingham (1852), Ipswich (1853), Worcester (1859), Lincoln (1872), Walsall (1874), and other places, some by commercial companies such as the Walsall Arboretum and Lake Company but most increasingly by town councils. The picturesque arboretum in Nottingham was noteworthy for its integration within a larger parks system, which was made possible by a large-scale enclosure act in 1845, which freed up common land for housing and park development. The scheme included a network of tree-lined avenues and parks. However, the botanical aspirations of these institutions as systematic tree collections tended to decline as their role as public pleasure gardens increased.\nAs one of Loudon's few realized park designs, much notice was taken of the Derby Arboretum. Downing, of course, had visited while on his tour in 1850. At the time, he was designing extensive public grounds in Washington, which incorporated a garden of American trees and a living 'museum' of evergreens, and he was actively urging the creation of a large park in New York. His experience observing British and European parks undoubtedly informed his thinking about the role of planting systematic collections. Although it was not executed, his plans for a public park in Boston for the Massachusetts Horticultural Society included a scientifically arranged arboretum.\nThe public parks of Britain provided important inspiration for Olmsted as well. Like Downing, he embarked on a tour of Britain, Ireland, and other parts of Europe in 1850. While he did not visit the Derby Arboretum on that trip, he made an inspirational stop at a new public garden in Birkenhead, a suburb of Liverpool. Like the Derby Arboretum, the gates of Birkenhead Park were open to the public without a fee 'but in this case for the whole week. It had been laid out by Joseph Paxton, who had designed other noteworthy landscapes including the arboretum and pinetum in the Chatsworth House gardens' Downing's favorite. Olmsted described Birkenhead Park as the 'People's Garden.' He was delighted by the winding paths and avenues and clusters of trees, set within wide, rolling lawns.\n'All this magnificent pleasure-ground is entirely, unreservedly, and for ever the people's own,' Olmsted wrote of Birkenhead Park. 'The poorest British peasant is as free to enjoy it in all its parts as the British queen.' The design and public function of Birkenhead Park would later serve as inspiration for Central Park. Olmsted revisited it as part of his investigation on the development of Central Park for the New York commissioners in 1859. On the same trip, he also paid a visit to the Derby Arboretum.\nA Public Arboretum in North American\nDespite growing interest in arboreta on both sides of the Atlantic in the mid-nineteenth century, a public arboretum with intentionally designed, labelled collections had yet to be established in the United States. There were proposals to plant the National Mall in Washington as an arboretum associated with the Smithsonian Institution, focusing upon American natives of some two thousand trees, and about two hundred species and varieties and counterpart to indoor natural history museum. Downing surveyed the landscape and produced designs for this in 1850 and 1851, after returning from his tour of British and European parks and arboreta. Support for concept of a national botanical garden had grown during the 1840s, including from Asa Gray, the professor of botany at Harvard and the director of the Harvard Botanic Garden. He had called for a national arboretum in 1844, emphasizing the research on American trees that had already been conducted by Andre and Francois Michaux and others.\nDowning's plan was for a public arboretum of labelled hardy trees and shrubs laid out in the natural style for educational and botanical purposes, and it included a pinetum. He also designed a picturesque garden surrounding the Smithsonian Institution formed with rare trees and shrubs. Although Downing's Washington plans were not implemented and Downing died in a boat accident in 1852 the concept of a national arboretum was ultimately realized outside the capital with the establishment of the Arnold Arboretum in 1872.\nThe Arnold Arboretum was integrated within a broader park scheme developed in Boston by Olmsted and the landscape architect Charles Eliot from the 1880s. The system, now known as the Emerald Necklace, consisted of a series of public parks connected by tree-lined parkways. Olmsted had proposed a similar concept in his report to the Brooklyn park commissioners in 1868. The integration of urban public parks using planted parkways hastened the development of urban forestry across the Atlantic world, and there was growing recognition that this was a distinctive endeavor which required special methods and expertise. There was also increasing emphasis upon the psychological and physical health benefits of trees in modern urban environments, although pollution, traffic, and buildings presented problems for planters.\nPart of Harvard University, the Arnold Arboretum would be free to the public all day, every day of the year. It expanded in a remarkably short space of time into a leading global arboretum guided by a director, Charles Sprague Sargent, whose longevity was hardly to be paralleled. However, the success also arose from its combination of elements of arboreta established across the Atlantic world over the previous century and collective body of arboricultural wisdom and experience. It combined picturesque naturalism with systematic tree collection, offering a place of study, recreation, and changing seasonal beauty. It was this that informed Sargent and Olmsted's collaborative design for the Arnold Arboretum.\nEgalitarian Ideals\nAlthough Loudon, Downing, and other arboretum promoters in the early and mid-nineteenth century argued that arboreta (like public parks generally) had recreational as well as scientific and horticultural functions, arboreta often remained associated with aristocratic and wealthy landowners and institutions with enough land, staff, and resources to form comprehensive collections with exotic trees and shrubs from around the world, some rare and expensive. The Arnold Arboretum's position as a part of Harvard University is a case in point.\nGiven these realities, nineteenth-century arboreta, like botanical gardens and parks, were idealized and often rather controlled, artificial, and regulated places. However, Loudon was motivated to promote them assiduously because he believed in their egalitarian possibilities, as did Olmsted. Loudon's gardening and natural-history magazines were intended to be forums that could be used by all social classes, from landed elites to gardeners, nurserymen, and women, and he strongly believed that gardeners ought to have a much fuller scientific professional education and have greater social status. As part of national, regional, or urban civic culture, arboreta had the power to transcend social divisions such as those between different social and ethnic groups (for example immigrant communities in North America) and between town and countryside, metropolis and nation.\nWhile nineteenth-century botanical gardens and arboreta were associated with trade, empire, and colonial exploitation, Loudon believed that this exchange of plant material would lead to global 'equalisation' of tree species, to the benefit all nations. 'If it is desirable for us that we should assemble in our country the trees and shrubs of every other similar climate,Loudon pointed out, 'it must be equally desirable that the inhabitants of every other similar climate should possess all those species for which their climate is adapted.'\nConclusion\nIn 1868, Josiah Hoopes, a nurseryman from West Chester, Pennsylvania, wrote that he believed his fellow American citizens were 'vastly behind' their 'transatlantic brethren' in the provision of tree collections' specifically collections of conifers. Yet, with the onset of the First World War, the initial decades of the twentieth century presented significant challenges for arboreta and gardens in Britain. Many of the arboreta established on country parks and estates declined because of general problems faced by the wealthy landed classes and their country houses after the war. British public arboreta such as those at Derby, Nottingham, Lincoln, Ipswich, and Walsall effectively ceased to be maintained as systematic tree collections for educational and scientific purposes and became indistinguishable from other urban parks.\nOn the other hand, with the professional development of forestry, urban forestry, and municipal horticulture, new arboreta were developed by the mid-twentieth century on both sides of the Atlantic. The most resilient British arboreta were those that remained parts of wealthy landed estates or academic institutions. Other long-term successes were arboreta that were acquired or developed by organizations such as the Forestry Commission or National Trust, the leading English quasi-governmental heritage organization founded in 1895 to 'preserve historical and natural places.'\nIn 1925, more than half a century after the creation of the Arnold Arboretum, Ernest Henry Wilson, the British plant explorer who became the Arnold's first keeper of the living collections, wrote that the number of visitors who journeyed from around the world to the tree collections in Boston increased by the thousands each year. He described the institution as 'America's Greatest Garden,' reasoning that because its raison d\u0550tre focused 'solely' upon the 'acclimatization, cultivation and study of hardy trees and shrubs,' the institution was entirely unique, even among European peers. Certainly, it had grown in a relatively short space of time into a peerless global institution, guided by Sargent, with a clear mission and supportive organizational structure.\nWhile Loudon's belief in the ideal of tree equalization across the continents is complicated in today's world of looming environmental crisis, the arboriculture practiced at the Arnold Arboretum from Sargent's day to the present has taken on a new urgency as the need to understand how trees respond to climate change becomes crucial. While Wilson's argument that the Arnold Arboretum brought man nearer unto man without boundary of race and creed remained an ideal rather than reality in an age of imperialism, oppression of Native American peoples, and continuing racial tensions, it is now beginning to be realized, aided by the collective desire to face the climate threat together as a global community, and to celebrate the symbolic value of public arboreta uniting trees from around the world for all to study and enjoy."},{"has_event_date":0,"type":"arnoldia","title":"Drawn to the Edges","article_sequence":9,"start_page":42,"end_page":47,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25784","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15ea726.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Angell, Bobbi","article_content":"Living in southern Vermont, I am surrounded by lush forests and verdant fields. There is so much to observe while trying to decide what to draw! Hobblebush (Viburnum lantanoides) has always been one of my favorite shrubs, with attractive winter buds, brilliant white flowers that light up the edge of woods in the early spring, and rich fruits in the fall, soon after eaten by birds. A single seedling, such as Pagoda dogwood (Cornus alternifolia) is far simpler to draw, but equally satisfying to find and turn into a copper etching.\nI have been drawing plants professionally for over forty years, and have always been obsessed with the precise detail that can be achieved with pen and ink. Several years ago, master printmaker Brian Cohen introduced me to the intricate art of copper etching, and I was immediately smitten. Accustomed to working within a defined space for publication, I appreciate the sharp boundaries of a copper plate. And since an etched copper plate is printed as a mirror image onto the paper, I work on my designs in reverse, checking it out on tracing paper and lightbox.\nThe final sketch is transferred onto a waxy ground application on a copper plate, and then I 'needle' it, scratching the wax with a sharp needle under my microscope, impressing fine lines and stipples, creating soft tones dot by dot. The plate is then etched with ferric chloric acid, printed as a test, and then reworked two, three, or more times to add more detail and depth. I use oil-based ink, rubbed onto the copper, cleaned first with fine cheesecloth-like fabric and then wiped with my hand. Each print is done, one by one, on water-soaked paper on my Ettan Press. I add watercolor to a select few of my editions. The editions are limited, usually 20 or 30 prints.\nPrimarily a scientific illustrator, I am attracted to unusual plants, reflecting my long history working with botanists and horticulturalists. I created a collection of such etchings for an exhibit with Beverly Duncan at The Arnold in 2018 (Impressions of Woody Plants: Disjunction, Two Artists and the Arnold Arboretum). As Beverly and I walked around the Arboretum with Michael Dosmann, planning the exhibit, I saw the gorgeous Chinese sweetbush (Calycanthus chinensis) in full bloom. Having learned it had been introduced into cultivation in the 1980s, I eagerly went out to purchase a shrub to grow, and draw, in my own garden. Also impressive is seven son flower (Heptacodium miconoides), the elegant, fall-blooming flowers and fruits of which I had seen at The New York Botanical Garden. Arnold Arboretum staff had collected seeds from a garden in China in 1980, raising plants for other institutions, including the NYBG. Within a few decades, seven son flower, too, had become commercially available, and so I was able to grow it and turn it into a copper etching. Tea viburnum (Viburnum setigerum), too, I first encountered at the NYBG, where I learned it had been introduced by the Arnold's E. H. Wilson in 1901. Corylus fargesii was commissioned for Curtis's Botanical Magazine, describing the introduction of a wonderful woody plant from China. The Arnold Arboretum has over a dozen plants in the collection. Magnolia loebneri 'Merrill', one of the finest hybrids ever released by the Arnold Arboretum, and named in honor of director Elmer Drew Merrill, was in full bloom during an early-spring visit to Smith College, stunning flowers displayed before the foliage leafed out. I was quite pleased when the Arnold Arboretum chose to use the resulting illustration for a logo for Arnold Selects (see page 7), a newly created program to bring exceptional plants from the living collections to gardeners around the world. "},{"has_event_date":0,"type":"arnoldia","title":"What in the World is a Species?","article_sequence":10,"start_page":48,"end_page":53,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25783","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d15ea36e.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Donoghue, Michael J.","article_content":"Many people are aware that species have formal names with two parts a genus name combined with what's called a specific epithet. Homo sapiens is a well-known example; for botanists, Ginkgo biloba will do. In their fullest form, they also include the name (or abbreviation) of the person or people who originally described the species. Homo sapiens was described by Carl Linnaeus in 1758, and in 1771 he named Ginkgo biloba, so you may see his initials after these names: Homo sapiens L., Ginkgo biloba L. There are very detailed (and ever-evolving) rules for how the description of a new species must be done for the name to be considered validly published. In botany, we refer to the International Code of Nomenclature for Algae, Fungi, and Plants for the exact procedures. It turns out that anyone yourself included can describe a new species if they follow these rules. You don't have to be certified as an authority to do this. Once you've published your new species, it generally would have one of two fates. Your new species could stand the test of time, in the sense that knowledgeable botanists would adopt it when they conduct their studies. However, unless you really know what you are doing, in 2022, it's likely you have named something that has previously been described. In this case, your proposed species name would be regarded as a synonym of the earlier one, and would henceforth be ignored.\nA key point is that you can validly publish a species name only to have it rejected by other botanists on the grounds that they don't consider it to be a 'real' species. This implies that there are some criteria being applied by scientists to judge whether something is a real species or not. It seems reasonable to assume that long ago there would have been agreement on what a species is on a species concept. This, however, is not the case. In fact, many different definitions of species have been published over the years, and to this day there are major camps of biologists who disagree (sometimes passionately) over which should be adopted as the universal standard.\nThe use of different species concepts by different scientists has a very important consequence: the various species that you are familiar with may not be equivalent to one another in ecological, evolutionary, or organismic terms. For the most part, however, we proceed as though they are. By 'we,' I mean not just the general public, but also the scientific community, who, despite knowing full well that multiple concepts are in use, still treat species as being somehow equal to one another. In reality, the only equivalence you can count on when you see species names is that they have been named according to some agreed-upon rules, and that they haven't been rejected by the scientific community. The potential non-comparability of species seems like a recipe for miscommunication. We proceed under the hope that species will somehow be 'equal enough' for most purposes, and that the differences among species won't interfere too much with scientific progress or public understanding.\nThe best-known definition, provided by ornithologist Ernst Mayr in 1942 and widely taught in introductory biology classes since the 1950s, is short and snappy: 'species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups.' This is the so-called 'biological species concept,' which many biologists accept in theory, although information on which organisms can interbreed is almost always lacking in practice. So, one generally just assumes such gene flow based on similarities and differences in the visible characteristics of the organisms, hoping that actual interbreeding will be tested directly someday. It has long been pointed out, however, that interbreeding and reproductive isolation aren't relevant criteria for organisms that reproduce through asexual reproduction. Such is the case with many bacteria, for instance, and with some plants as well. And there's the associated question of whether any level of interbreeding could or should be tolerated. This has been a special concern for botanists, where hybridization is often possible between species that appear to be quite distantly related (consider all of the strange orchids that have been produced in this way).\nAlthough the biological species concept is the most widely known, there are a variety of alternatives that feature different criteria. One such alternative focuses on species as occupying particular ecological niches that differ from related species. Another one focuses on shared common ancestry, delimiting species based on evidence that certain organisms and populations share a common ancestor separate from related species.\nOne concept I find especially appealing is known as the 'evolutionary species concept,'proposed by the paleontologist George Gaylord Simpson in 1951. Working with fossils of long-dead mammals, he wanted to take the emphasis off of interbreeding (which he certainly couldn't test). Instead, he conceptualized species in terms of a full evolutionary life cycle, from inception to extinction. Simpson said a species is: 'a phyletic lineage (ancestral-descendant sequence of interbreeding populations) evolving independently of others, with its own separate and unitary evolutionary role and tendencies.' Under this view, the populations that we study today are time slices through an extended lineage evolving independently of other lineages. This concept provides a nice image of species, though for many people, 'role and tendencies' have seemed a bit squishy and difficult criteria to apply in practice.\nOne very nice 'solution' to the species problem was proposed by herpetologist Kevin de Queiroz in 1998, and reinforced in his subsequent work (e.g., de Queiroz, 2005). He noted that all of these concepts focus on populations or lineages extended through time and evolving independently of one another. In his view, reproductive isolation, ecological differentiation, and exclusive shared ancestry may arise in different temporal sequences as the process of speciation (the origin of independently-evolving lineages) proceeds. At any given point in the process, species might have some of these properties, and not others. For example, gene flow may be cut off early in the process, perhaps by the simple geographic separation of populations, as compared to, for example, ecological differentiation.\nUnder de Queiroz's so-called 'general lineage concept' of species, phenomena formally viewed as necessary and sufficient defining criteria for species-hood, are instead understood to bear on whether, in fact, two lineages are evolving separately. If we find, for example, that the organisms in two populations are unable to breed successfully with one another, this provides pretty good evidence that the populations are evolving separately. Likewise, the finding that populations are occupying different ecological niches provides evidence of independence, as do consistent differences in morphological characteristics. These things don't define species, but instead help us to discover them.\nThe general lineage concept of species has been steadily gaining popularity among evolutionary biologists, but it is still far from universally accepted. Personally, I like it very much, but would stress a few additional points. First, I think that the delimitation of a species is best viewed as putting forward a hypothesis to be tested with evidence of lineage independence coming from as many different angles as possible. By this I mean to include not only information on breeding, but on geography, morphology, DNA sequences, ecology, and a host of other criteria. Second, I would like to preserve Simpson's reference to the future and predicting the likely fate of a lineage. It seems reasonable to add into the decision-making process whether it seems likely that two lineages will continue to evolve independently into the future. Evidence bearing on fate may also come in different forms. For example, consider the two species of tulip tree: the familiar eastern North American Liriodendron tulipifera, and the eastern Asian Liriodendron chinense. These can readily be hybridized, and the offspring plants (L. tulipifera chinense) are fertile. Living proof of this can be found at the Arnold Arboretum, on the lawn in front of the Hunnewell Building. But, it seems reasonable to suppose, based on their very widely separated geographic ranges, that individuals of these two species will not naturally be exchanging genes any time in the foreseeable future. Finally, I also really like the reference to 'tendencies,' as this highlights the idea that a separately evolving lineage will often show a propensity to generate certain variants again and again as compared to another species. Mind you, I don't at all mean to suggest that such tendencies should define species; rather, in keeping with the general lineage concept, they can potentially serve as evidence of independent evolution.\nAllow me to end with a few observations about my own favorite plant group, Viburnum. When I was a graduate student at Harvard, in the late 1970s, I lived on the grounds of the Arnold Arboretum, at what used to be 383 South Street. Of course, I wandered the grounds often, and it was there that I became well acquainted with around 40 of the roughly 165 Viburnum species, many of them from eastern Asia, where Viburnum is the most diverse. You can learn a lot about species differences in an arboretum, but not nearly enough to critically assess their evolutionary independence from one another. For one thing, you don't see the species that can't be grown in the arboretum (e.g., Viburnum species from tropical forests in Borneo, or from high elevations in the Andes), or the many species that could potentially be grown but have never been brought into cultivation. And, you really need to study organisms in their natural surroundings to understand the range of variation that they exhibit, their ecological niches, and which species might encounter one another in the wild.\nI did, however, manage to observe something about Viburnum species that has turned out to be more important than I ever imagined. I went out on a regular basis to record the times when plants of different Viburnum species were flowering in the arboretum. I found that they were flowering each year in a consistent sequence, staggered through the spring and early summer. In fact, these observations were the basis of my very first publication, in 1980, which happened to be in Arnoldia, and was entitled 'Flowering times in Viburnum.'\nAs we have learned since that time, related species of Viburnum living in the same geographic area very often flower at different times, which means that they are reproductively isolated from one another in this temporal way. For example, as shown recently by my former graduate student Elizabeth Spriggs, the species of the Viburnum lentago complex in eastern North America (nannyberry and its relatives) bloom at different times, and this minimizes hybridization between them where their geographic ranges overlap (Spriggs et al., 2019a; Spriggs, 2019). We know that individuals of these different species can breed together successfully. In fact, Viburnum jackii, a hybrid between V. lentago and V. prunifolium, was described from a plant first noticed in 1908 at the Arnold Arboretum. However, in the wild these species rarely do hybridize, simply because they are flowering a week or so apart. Importantly, given the discussion above, I am not supporting the biological species concept with this observation. Instead, I am adopting the general lineage concept and using this flowering offset as one line of evidence that these are time-extended lineages evolving on their own.\nI hope that these few reflections will heighten your appreciation of species when you see your next specimen label in the Arnold Arboretum-- perhaps even a Viburnum lentago L. plant in the superb Viburnum collection near the Centre Street Gate!\nReferences\nde Queiroz, K. 1998. The general lineage concept of species, species criteria, and the process of speciation. In D. J.\nHoward and S. H. Berlocher, eds. Endless Forms: Species and Speciation. Oxford University Press. Pp. 57\u00d075.\nde Queiroz, K. 2005. Ernst Mayr and the modern concept of species. Proc. Natl. Acad. Sci. USA 102: 6600\u00d06607.\nDonoghue, M. J. 1980. Flowering times in Viburnum. Arnoldia 40: 2\u00d022.\nMayr, E. 1942. Systematics and the Origin of Species. Columbia Univ. Press, New York.\nSimpson, G. G. 1951. The species concept. Evolution 5: 285\u00d0298.\nSpriggs, E. L. 2019. The Viburnum lentago clade: A continental radiation. Arnoldia 77: 10\u00d019.\nSpriggs, E. L., C. Schlutius, D. A. R. Eaton, B. Park, P. W. Sweeney, E. J. Edwards, and M. J. Donoghue. 2019a. Differences in flowering time maintain species boundaries in a continental radiation of Viburnum. Amer. J. Bot. 106: 833\u00d0849.\nSpriggs, E. L., D. A. R. Eaton, P. W. Sweeney, C. Schlutius, E. J. Edwards, and M. J. Donoghue. 2019b. Restriction-site-associated DNA sequencing reveals a cryptic Viburnum species on the North American coastal plain. Syst. Biol. 68: 187\u00d0203."},{"has_event_date":0,"type":"arnoldia","title":"A New Way for the Norway Mapl","article_sequence":11,"start_page":54,"end_page":57,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25782","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d14e896b.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Contreras, Ryan","article_content":"The summers of my youth in Eastern North Carolina smelled of Chinese privet (Ligustrum sinense) and Japanese honeysuckle (Lonicera japonica). As a kid, I loved playing with the tiny 'berries' of the privet and sucking the nectar from the honeysuckle flowers. Warm memories aside, these two species are landscape plants turned weeds, which escaped cultivation and invaded large areas across the Southeast. As someone who works with the nursery industry and specifically with this issue of weedy or invasive plants, it sometimes feels that folks believe all introduced plants are bad, and we should only grow natives to protect our ecosystems.\nWe should think, however, about what is it we are asking our landscape plants to do. In the city, we want them to survive stress, even to flourish. We want to punish them with drought, heat, pavement, and poor and compacted soils while still enjoying their shade, beautiful flowers, lovely scent, and fruit. Whether native or introduced, plants that thrive well enough to escape cultivation are doing exactly what we asked of them.\nI often hear that we should only plant native plants because they are best adapted to a site or region. If that is the case, how do the non-native and introduced species outcompete them? There also are 'native' plants that have become 'invasive': western juniper, for instance, now covers more than 2 million acres of grassland in Oregon, its spread aided by fire suppression. We need plants that do well in our cities. We should care less about their provenance and focus more on their behavior. The problem isn't trees that flourish, but trees that won't stay where we put them.\nTake the Amur and Norway maples, two resilient species commonly found in our cities. Easy for producers to grow, they thrive where other species may not survive. Amur maple is hardy to USDA Zone 2, fitting the bill for a small urban tree in regions short on options of plants from which to choose. Norway maple is hardy to USDA Zone 4, making it suitable as a medium to large tree in most of the US. Both are relatively free of major pest problems, and transplant well. Norway maple is also incredibly well-adapted to heavy clay and compacted soils, tolerates pollution, and holds up better to drought conditions than sugar maple. Unfortunately, both have done their job too well, and have escaped cultivation to invade native forests and cause real problems in several parts of the country. As an urban tree, however, they fit the bill incredibly well, helping to ameliorate the heat-island effect, manage stormwater, and beautify our paved metropolises. It is not surprising that such resilient trees can outcompete other species.\nOn Burnside Avenue in downtown Portland, Oregon, just down the street from Powell's City of Books, there is a planting of Norway maple that separates opposing lanes of traffic. The soil volume is tiny, and tall buildings loom on either side. Yet, these Norway maples are gorgeous; more than 35 feet tall and healthy, they cover most of the five-lane driving surface and cast shade on the sidewalks for pedestrians. Contrast this to urban instances of our native bigleaf maple, such as the large specimen near Valley Library here on the Corvallis Campus, or the majestic tree that greets you as you set out on the trail at Hoyt Arboretum in Portland. These are 'easy' sites for trees, with large soil volumes and little compaction. You will not find bigleaf maples adorning streets like Burnside Avenue, however.\nWe could alter conditions to suit bigleaf maple redeveloping our cities for more soil volume, less concrete, and less pollution but that does not seem likely. Alternatively, we could breed more resilient bigleaf maples a path that is being explored, but likely will take a very long time.\nMy research program is making great progress pursuing a third option: breeding Amur and Norway maples that stay put where we plant them. We want to provide growers, land managers, and the public the utility of resilient trees that are good for cities, but also do not reproduce in sufficient numbers to displace our native flora.\nHere, it's worth mentioning 'Bradford' pear. Perhaps the most numerous of the many cultivars of Pyrus calleryana, it has become the poster child for invasive plants. Smelly, weedy, fragile in ice storms, it's the tree people love to hate. 'Bradford' and other pears are self-incompatible, which means they need another genotype to fertilize their ovules and form seeds. Soon as new cultivars were introduced these genotypes started cross-pollinating and producing fruit, soon becoming the weed we know today. Pyrus 'NCPX2', the Chastity\u00a8 pear developed by Tom Ranney of North Carolina State University, by contrast, was recently tested for fertility compared to wild-type, and is not merely self-incompatible. Chastity\u00a8 is a triploid that is, it has three sets of chromosomes. This odd ploidy (number of chromosome sets) disrupts normal formation of pollen and eggs, resulting in a plant that infrequently or never produces viable seeds. The most famous triploid out there is banana. If you have enjoyed a 'Cavendish' dessert banana, then you have enjoyed a delicious fruit rendered seedless through triploidy.\nThough there are reported examples of Norway maple exhibiting reduced seed set or seed germination, in my experience these cultivars are perfectly fertile. It is unclear in what contexts the trees have set seed, but these cultivars are not sterile across environments thus my reluctance to use the word 'sterile' in context of seed set. As with most cases in nature, there is a gradient from perfectly fertile wild-type down to complete sterility. As such, I try to stick with 'reduced fertility' as the descriptor for cultivars that reproduce at such a low level as to pose no ecological threat.\nThe first step in the process was to induce chromosome doubling of standard diploid plants (containing two sets of chromosomes) to develop tetraploids (plants with four sets of chromosomes). We planted our tetraploids alongside diploid cultivars at our field in Corvallis and allowed them to open pollinate. We collected seed from the tetraploids, grew seedlings, and tested their ploidy level. Fortunately, most of these seedlings were triploid they received two sets of chromosomes from their tetraploid female parent and one set from their diploid male parent. Furthermore, these seedlings are not genetic composite (chimeras), but are triploid in all cell layers, and thus highly stable from one generation to the next.\nTen years after starting this project, I published the results of this work in 2020 in the journal Horticulturae. But the work in so many ways is just beginning. To produce the seedless trees we desire, they must be propagated clonally. Traditionally, Norway maples (and Amur, too, in some nurseries) have been chip budded, grafting the cultivar of interest to seedling rootstocks. While this production system speeds up the production and quantities of triploid clones, we need a new tactic. This is because we must avoid at all costs grafting our sterile triploids onto fertile diploid rootstocks rootstocks that can sometimes send up their own shoots and eventually produce seeds, which happened with callery pear. For Amur maple, this is not a major problem, as it readily roots from stem cuttings. For Norway maple, which does not, we have been working to optimize cutting propagation. We now have triploid genotypes of both species, which we are growing via micropropagation, using sterile culture in vitro to multiply plants in large numbers relatively quickly. This technique is used in many taxa that would otherwise be slow to increase using other methods, such as hazelnuts (Corylus avellana). It also is frequently used in red maple as a means to increase and distribute clean clonal material. Our triploid plants will be ready to come out of micropropagation and harden off to begin production trials during 2022.\nEvidence of reduced fertility gives us much reason to hope. Amur maple triploids in our plots have flowered in the presence of pollinators and fertile pollen donors and have produced no viable seeds to date. While this inspires confidence, I am not ready to bet the farm or rather, to have growers bet theirs. Our next step is to work with nurseries, universities, and public gardens around the country to install replicated tests of our trees to see how they perform in other environments. The stakes are too high not to verify.\nOf course, my title is Ornamental Plant Breeder, so the trees resulting from this work should have some aesthetic appeal. To that end, we are working with J. Frank Schmidt and Son Nursery along with Tom Ranney to evaluate seedlings of Amur maple selected at JFS in Boring, NC State in Mills River, and Corvallis, OR. Ten genotypes from each location were propagated during 2021 under production conditions to identify superior forms. Furthermore, the trees from micropropagation will be included in a parallel study. The end goal is to develop and test trees according to the best scientific methods we have, while working with growers to ensure that we are meeting their needs for trees that work in production.\nThere is no doubt of the need. Industry partners report more than 90% reduction in Norway maple sales, with steep declines in Amur maple as well. Certainly, overplanting of maples has reduced demand, but the invasive issue has also had an impact, and the industry is ready for cultivars of these species that could be sold in longstanding markets such as the upper Midwest and New England.\nEvidence indicates the trees I have developed (and those of my colleagues like Dr. Ranney) are 'sterile,' or close enough that they present no threat of invasion. The biological side of the problem is largely solved. What remains is the political aspect, which in many ways is more difficult. The story of 'Bradford,' damaging in its lack of nuance, has spread effectively, and plants like Norway maple may prove difficult to reintroduce as a result. Already it is illegal to plant A. platanoides in Massachusetts, and many other states a rule which leaves no room for reduced-fertility cultivar exemptions.\nWe need a national conversation on this topic in the Green Industry, to collectively establish the framework for reintroduction of sterile versions of weedy species. The specifics of individual plants are highly regional, and thresholds should be determined at a state level, but the issue is a national one. The shade of that tree you're enjoying on the east coast may have gotten its start here in Oregon. As such, the rules enacted in Massachusetts have wide-ranging impact. The need for education, collaboration, and nuanced regulation will only grow, so long as cities remain, and climate change increases the demand for resilient trees."},{"has_event_date":0,"type":"arnoldia","title":"What Clings to the Roots","article_sequence":12,"start_page":58,"end_page":59,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25781","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d14e8927.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Battles, Matthew","article_content":"The morning we moved out of my childhood home, the new owner pulled up with a small tractor to uproot the Forsythia hedge, my mother's pride. It was late April, I think, as the bushes were in bloom; sprays of yellow blossoms shivered as the backhoe groaned and clawed at the plantings. I was shocked by how easily they came up, ungainly roots whipsawing as they shook loose from earth. My mother sobbed as we drove away. And yet soil clings to the roots; an ecology shifts intact. To uproot is an ambivalent move, metaphorically: is it about the fragility of attachments, or their stubbornness to endure?\nSalom\u017d Jashi's Taming the Garden opens with a tree shimmering on the horizon, rooted in the liquid tumble of the sea. Lashed to the deck of a barge, its headway is barely perceptible against the lowering sky. The barge sails under the orders of Georgian oligarch Bidzina Ivanishvili, whose minions search farm and forest for the prodigious trees he has uprooted and moved to his 'dendrological park' in Shekvetili, a resort town on Black Sea coast. We never see Ivanishvili; no agents or officials sit to offer apologies or explanations to the camera. The oligarch's name is only occasionally uttered by workmen and townspeople, and he remains a minor character, his motives a mystery to the people whose trees he takes. One man claims to have read that 'it prolongs his life' to collect trees, if their age is greater than one hundred years. Some praise his enterprise, while others boggle at the cost of the operation. 'No matter how much a villain he is,' another exclaims, 'at least he's doing something!'\nTownspeople gawk at their trees on the move. Their faces register the dappled play of emotions, from grief to wonder, as workers cut, dig, and lever at giant trees a towering tulip, goblet-shaped and elegant; a bounteous linden growing close by an old house; a chestnut with two splayed leaders that swing like the arms of a drunken giant. Their slow severance from the earth is both clumsy and precise, a kind of terrestrial surgery, at once an amputation and a deliverance of tender care. Jashi allows the sensuous overwhelm of these labors to fill her frame: a trench dug round the tree, the earth wrapped with sheets and shored up with boards, and a framework of pipes bored through below, driven home with rust-streaked drilling augers. The scale of the work matters to Jashi: we see men chopping, sawing, dragging brush, dwarfed by walls and mounds of foliage. A backhoe swings into view, framing the shot like a great mechanized tree; from another angle, viewed downslope through a colonnade of what look like hemlocks, the same machine looks minuscule. During a break, the crew sit around a fire of brush and reminisce. They agree that the trees are very beautiful. 'Life takes strange turns,' says one.\nJashi is a generous storyteller, and patient. Long takes invite us to ponder how a mature tree organizes its surrounding space: the way the earth bunches muscularly at the roots; how its shade selects and prunes the vegetation; above all, the way it pigments and concentrates the air in its branches. And then we watch the slow, uncanny spectacle of this composition deconstructed, as yet another great tree is carved out of the ground, jacked onto a carriage, and towed off, leaving a crumbling pit of soil to fill up with new vegetation.\nI think of those islands of earth cut and carried away, with their cryptic assemblages of fungi and invertebrates, to be installed in the oligarch's faraway estate, ferns and flowering plants bobbing in the shade of a tree transported over the sea. The trees' communities exceed grasses, forbs, and fungi, however, rooted as they are in the loam of family and village. Local people gather in the night to watch as a towed tree sways in spotlight gleam. 'It's so beautiful in the night,' one says. 'Like a fairytale.' 'It won't survive,' says another, 'it's shrunk so much.' An old woman confronts the cutters: 'she planted this tree,' her companion warns the foreman; 'what we do in this world will be judged in the next.' Elders embrace, young people shoot video on cellphones, the tree moving stately through pines as flashlights lance through the galleries of boughs, the lights of the trucks closing in, filling the frame, branches of roadside trees snapping as the tulip shoulders through. Jashi stays with these shots a long time, lingering in the strangeness of a tree swaying in the still of night.\nWhat are we to make of Ivanishvili's uprootings? How do we weigh the ecological and social costs; how does his project compare to the collecting practices of public gardens and arboreta? Jashi eschews such ready questions and contrasts, preferring to dwell patiently in the confusion of the more-than-human encounter. Resisting easy critique, her eye is anthropological, tracing the exertions of people and trees with equanimity and affection. Along with townspeople and workers, we're invited to boggle, mourn, and wonder. And the trees in the end are beautiful, settled in their new home amid sprinklers and curving paths.\nIn his lavish account of Kublai Khan's pleasure palace, Marco Polo describes a hill planted with mature evergreens collected throughout the empire and carried to the capital by elephants. Historically, Marco Polo arrives on the eve of modernity and the coming Anthropocene before the forests of North America travelled upright over the seas in the form of ships' masts; before the forests of Asia and South America were felled for tea and palm and rapeseed. The ecological impact of Ivanishvili's Dendrological Park pales by comparison to such depredations. It's even beautiful in its way. Like the green hill of the Khan, the oligarch's park is lush, verdant, well tended. The birdsong there is fluting and evocative. And yet the trees are still rigged with the cables, bound fast like wild beasts. The oligarch wants his country to behave like a well-loved garden. And yet, as Salom\u017d Jashi reminds us, the memory of living soil persists amid the roots."},{"has_event_date":0,"type":"arnoldia","title":"Of Trees and the City","article_sequence":13,"start_page":60,"end_page":62,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25780","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d14e856f.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Stephens, Matthew","article_content":"Anyone who has planted and cared for a new tree knows that few things in life are as rewarding as this simple act. Planting day is unquestionably a stand back and be proud moment, but those of you who have planted one tree, or many, know that the real work (and appreciation) is just beginning. The watering, weeding, pruning, and care that is needed is an investment that will pay back dividends in seeing a tree grow bigger and bigger with each passing season.\nI happen to fall into a special category of tree planters: someone who can take credit for having played a significant role in planting over one million trees. This is becoming less of an incredible accomplishment given many places are now planting millions or billions of trees to combat climate change. However, there are few who can claim such a large bounty in an urban area, and specifically New York City. Prior to becoming the current President & CEO at Mount Auburn Cemetery, I helped to lead MillionTreesNYC, an effort to plant one million trees throughout all of New York City between 2007 and 2015. We planted trees along streets, in parks, and in cemeteries and botanical gardens work that will continue indefinitely, just as occurs in nature.\nOne question I have been asked about my work is, 'how do you plant a million trees in New York City?' The answer can be reduced to a simple instruction: one tree at a time! Further, New York City should be applauded for its efforts to use MillionTreesNYC as a springboard to further investment in pruning, permitting enforcement, and staffing tied to managing the urban forest.\nHaving landed at Mount Auburn in September 2021 as its new President & CEO, I was immediately entranced by the awe-inspiring collection of oaks and beeches. Without question, Mount Auburn has one of the best collections of mature trees anywhere in the United States, with nearly 4,700 trees of varying ages and over 650 taxa on its 175 acres. During a ten-minute walk on the grounds, you are sure to see specimens of multiple species of trees that will be some of the best you'll ever see! Our trees, some over 200 years old, have seen the world reinvent itself many times over, yet continue to reach for the skies with each passing year. For nearly 200 years our trees have received remarkable care in the form of watering, pruning, and other conscientious landscape maintenance techniques which have allowed them to thrive. Further, trees at Mount Auburn don't have the same competition as most urban trees.\nWhile tree planting traditionally gets the most fanfare and showy pictures, the years of effort and care leading up to a canopy-covered street tend to be overlooked. A few steps beyond our gates I am reminded of how tough it is to be an urban tree, especially a street tree. Between traffic, dogs, developers, climate change, and countless other variables, these trees face many stresses which shorten their lifespans. Struggling to keep up with necessary tree maintenance, cities worldwide have backed away from tree planting goals while also minting goals for canopy coverage. Ultimately, it is every urban forester's hope to invest resources, create policies, and develop stewardship to increase the canopy percentage over time.\nTrees happen to be quiet constituents. Rarely will an email, phone call, or press conference intervene when a community tree is suffering, unless an urban Lorax intervenes. Trees take time to grow; a future canopy doesn't develop on the schedule of politics and budget cycles. A tree planted today will take decades to equal the annual ecosystem services generated by the biggest and most beloved trees. This is a tough reality for trees in all our communities. However, the data that have been collected over the last three decades enunciate with extreme clarity: mature trees, and especially large shade trees, are exponentially much more significant providers of the ecosystem services. The math is simple: the larger the tree and more leaf surface area, the larger the benefits. For example, a newly planted tree, just a few inches in diameter, may sequester six pounds of carbon, or currently valued at about thirty cents; a mature tree greater than thirty inches in diameter, by contrast, will sequester over 6,000 pounds of carbon, worth some four hundred dollars. A thousand-fold increase! With that, how can trees continue to be overlooked?\nBeyond their value as carbon store, trees provide real and tangible benefits in the form of cleaner air, shade for buildings, or stormwater capture among many, many others. Many years ago I remember talking to Dr. David Novak with the US Forest Service who has dedicated his career to studying urban forests. Comparing the urban forest to other forms of infrastructure, he mentioned that we are just starting to fully realize the benefits of trees. Walk down the street where you live, and you will see some permutation of city infrastructure: fire hydrants to ensure buildings don't burn down, light poles to provide safety, or stop lights to allow traffic to be regulated, among others. Funded through local, state, or federal dollars, these investments improve the quality of life or safety of a given neighborhood. Compared to trees, however, light poles have lower dollar value in benefits and unlike trees, they decrease in value over time.\nWhy, then, have trees gone so overlooked as critical parts of urban infrastructure? Simple: trees are rarely considered a capital investment. But, if they were, it would provide urban foresters access to new and necessary sources of funding. Additional funding and pragmatic, focused local tree preservation legislation are long overdue. Trees should be funded, along with highly competent urban forestry managers to manage the urban forest which, like all critical urban infrastructure, is key to the safety and well-being of residents. In addition, many cities have a mechanism in place to raise capital monies through the selling of municipal bonds why couldn't trees be included along with other key infrastructure that elevates the quality of life of a locality?\nMany cities are making great strides, but there is still much work to be done. During my time in New York City, I would travel the country helping other cities figure out how to attract more funding for trees. Some cities were incredibly creative, but a clear thread emerged: urban forestry managers must scratch and claw for every dollar they get. And trees get pennies on the dollar compared to other urban infrastructure. In many cities, public\/private partnerships are aiming to fill the gaps. From Washington, DC, to San Francisco, to Portland, robust and sophisticated urban forestry nonprofits are filling the gaps left by public funding.\nOne irony of this struggle is that many cities or towns have left tree management\/urban forestry to a roads and sidewalks or public works department the areas of government that typically manage infrastructure. As a result, urban forestry programs have modest resources and\/or no meaningful political support given they are buried in large public works departments, and must compete against potholes or sidewalks for attention and funding. The reality, however, is that a well-sited tree likely will outlive all its infrastructure counterparts, outlasting sidewalks, stoplights, and even many buildings.\nFurther, local tree legislation that protects trees on public and private property is also lagging. Every city desires some level of development; however, it has also been the experience of many urban foresters that the impacts trees encounter from new construction, sidewalk\/driveway work, or other infrastructure projects lead to a significant number of removals or tree mortality after construction is completed. While a tree may not die immediately from construction impacts, my time working in New York suggests trees must be monitored for several years post construction to fully assess development impacts. When I met with developers in New York, they were quick to point out that they will likely spend more on doorknobs or cabinet handles than they will on trees even though the trees become part of infrastructure, and a community asset. When replacement is mandated by local legislation, it often merely requires a 1:1 planting ratio such that an old mature oak tree in its prime, for example, might be replaced with a newly planted red maple. We know from the data, however, that a newly planted tree can't replace a fully-grown tree in the urban infrastructure. There are few cities like New York City who are using a basal-area replacement methodology, which is a more appropriate way of calculating the true cost of removing healthy trees. That calculation not only more adequately accounts for loss, but protects trees by ensuring that any developer thinks twice before removing a tree.\nThe time for policy change is now. We need those who will speak for the trees, knowing they are a critical part of the urban infrastructure. Find fellow Loraxes, and organize. Approach your local elected officials and let them know how important the trees are to you and your community. Work with them to move forward thoughtful and pragmatic legislation. It will take time, steadfastness, and collective action by like-minded citizens who can speak and act civilly and passionately to make change change that will, that must, happen one tree at a time.\nIf you are in the Boston area, I encourage you to stop by Mount Auburn to check out our incredible canopy in a thriving metropolis. I guarantee you will leave feeling inspired by our one-of-a-kind landscape. Then, find a tree in your own neighborhood and start giving it some care. I am certain the time and energy you invest will be repaid in dividends. Enjoy your trees! "},{"has_event_date":0,"type":"arnoldia","title":"Deadheading Lilacs","article_sequence":14,"start_page":64,"end_page":64,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25779","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d14e816c.jpg","volume":79,"issue_number":"2","year":2022,"series":null,"season":null,"authors":"Guidarelli, Connor","article_content":"The Lilac Collection has been getting its very own special day of celebration, Lilac Sunday, every Mother's Day for the past 112 years. Flowering extends beyond this day, of course, running from the end of April to the beginning of June. Within 2 weeks of flower wilt, we begin preparation for next year's spectacle by deadheading the lilacs. This practice helps to ensure that the shrubs do not expend more energy in seed production, but rather use it to produce flower buds more prolifically.\nMany hands make quick work of this time-sensitive task, as interns, seasonal gardeners, and horticulturists make their way through over one hundred plants. Some shrubs are so large that we need our six-foot extendable pruners to reach many of the spent flowers. Orchard ladders extend our reach even further, making it easy to maneuver in and around a shrub. Between plants, we spray sterilizing solution on our snips to prevent the spread of pathogens like phytoplasmas, often called Lilac Yellows. All the cuttings are collected and composted, to return to the collection as a soil amendment come the fall."},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25778","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d14e8128.jpg","title":"2022-79-2","volume":79,"issue_number":"2","year":2022,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Arnold at 150","article_sequence":1,"start_page":1,"end_page":4,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25755","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170ab6e.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Friedman, Ned","article_content":"What is the Arnold Arboretum? This question has been at the center of my thinking for over a decade, especially now, as I enter the twelfth year of my directorship and the Arnold enters its 150th year. Of course, nothing should ever be static when it comes to the life of an institution. Founding nineteenth-century ideals need updating in the twenty-first century. Still, for all that has changed over the last century and a half, the core values of the Arnold Arboretum strike me as eternal.\nThe Arnold Arboretum has and will always serve as a crossroads for biodiversity and human diversity. Its founding was a testament to the enduring values of democratic spaces (free and open to all) and the belief that such places should uplift all who enter. The Arnold is also, from the outset, an institution defined by its association with Harvard University. Scholarship, born of a love of biodiversity and a desire to unlock its secrets, is central. An ethos of conservation and respect for the environment goes back to the founders and early leaders. The meanings of such an intermingling of sentient and nonsentient organisms (respectively, people and trees) can never be fully unpacked, even in a lifetime of pondering. Yet I will briefly reflect on my thinking.\nLet's begin with my definition of an arboretum: a collection of woody plants with provenance in a designed landscape. Here, provenance and designed landscape are essential characteristics that help us appreciate the varied and dynamic relationships that occur between people, uniquely identified botanical organisms, and arboretum landscapes. The concept of provenance is typically associated with museum objects (think artworks), and at the Arnold Arboretum, every organism has a documented and acknowledged history. Take, for example, a single specimen of the sand pear (Pyrus pyrifolia, accession 7272*C) that has grown on the top of Bussey Hill for over a century.\nWe know that Ernest Henry Wilson and his collecting team encountered the parent of this sand pear growing west of Yichang, China, in the late summer of 1907. They collected fruit, removed its pulp (perhaps by eating it?), and separated, dried, and packed the seeds. The packet then passed as cargo down the Yangtze River to Shanghai, made its way by steamer to the west coast of North America, and took the transcontinental trains to Boston. On April 15, 1908, an Arboretum propagator formally accessioned the seeds. A few years later, a spot for a young sapling was chosen, and a hole was dug. This wonderful organism has lived in this location ever since, battling plant diseases and delighting visitors with its extraordinary clouds of white flowers every spring. This specimen is not any sand pear. It is an individual with its own life history and standing, not interchangeable with any other sand pear on Earth, just as no two human beings are interchangeable. Such provenance granular and unique distinguishes almost all the Arboretum's roughly sixteen thousand accessioned woody plants.\nA designed landscape is also central to my definition of an arboretum, and the Arnold Arboretum is fortunate to have been designed by a visionary Frederick Law Olmsted. His intentional design is reflected in every inch of the grounds, like the majestic reveal as you round the bend on Hemlock Hill Road and unexpectedly view the dramatic mixture of spruces and firs, with their blues and seemingly endless hues of green. The intentionality can be felt as you stand under the cathedral-like oak collection or take in a seemingly endless run of mountain laurels in flower in the spring. This landscape was designed to affect us and, indeed, to lift our spirits every day.\nThe impact of these experiences is profound. Olmsted spoke of the power of institutions like the Arnold Arboretum 'to make life in the city healthier and happier. But, surely Olmsted, despite his public health credentials (as general secretary of the US Sanitary Commission during the Civil War), would never have dreamed of the slew of well-documented health benefits of beautiful urban green spaces such as the Arnold Arboretum. Those who regularly walk these grounds may experience (on average) lower blood pressure, improved postoperative recovery, improved birth outcomes, improved outcomes associated with congestive heart failure, improved child development, reduced mortality, reduced stress, reduced symptoms of attention deficit hyperactivity disorder, reduced depression, and greater life satisfaction the list goes on. The Arnold Arboretum is literally interwoven into the healthcare system of Boston.\nOn a global scale, the research and conservation functions of the Arnold Arboretum have never been more critical. Fully three-quarters of the research now being conducted in the living collections is centered on understanding and combating human-induced global change, including climate change. How will trees and forested ecosystems function going forward, as climactic extremes mount by the year and invasive pests and pathogens circle the globe? The Arnold's working collection of woody plants is on the job providing essential insights into the coming biological Armageddon. Our plant expeditions throughout the temperate regions of the Northern Hemisphere emphasize the collection of germplasm from species and populations that are threatened with extinction. Ex situ conservation, the maintenance of living collections of endangered plants in botanical gardens and arboreta, has never been more critical to the Arnold's mission and to Earth's botanical biodiversity.\nI could go on but will finish by reflecting on the last two years of the Arnold Arboretum's existence. Through a raging and lethal pandemic, a reckoning over systemic racial injustice, an insurrection and serious challenge to American democracy, and the ever-more obvious extreme fires, floods, droughts, heat waves, and other threats to the world's four-billion-year evolution, the Arnold Arboretum did not close for a minute.\nThe Arnold Arboretum is not a mere amenity or simply a pleasure ground. It is an essential part of the public healthcare system, a place where the diverse population of Boston mixes, a bulwark for democracy, a leader in fighting global change and extinction, and a place where the next generation of ecologists, evolutionary biologists, and conservationists will launch their careers. And standing behind all of this are the magnificent plants with provenance in an Olmsted-designed landscape. What could possibly be more beautiful and meaningful as the Arnold Arboretum launches into its next century and a half?"},{"has_event_date":0,"type":"arnoldia","title":"Planting the New Lions of Kew","article_sequence":2,"start_page":8,"end_page":8,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25760","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170b76a.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Kirkham, Tony","article_content":"As head of the arboretum at the Royal Botanic Gardens, Kew, I would walk the collection each day, choosing a different route and corner of the three-hundred-acre landscape. On the walks, I observed the growth of newly planted trees and built up a knowledge of the collection. I wanted to understand where gaps occurred and what we should plant to improve the wealth and diversity of the woody collections. I kept an eye out for important but ailing plants that should be repropagated. This daily practice remained valuable no matter how long I worked at Kew a tenure that spanned forty-three years in various roles.\nI describe the arboretum at Kew as a living reference library of woody plants from every corner of the temperate world that will grow outdoors (near London) without any form of protection during the winter. However, overseeing a collection like this isn't just about planting trees as they become available and looking after them. The collection is visited by two million people per year. It must meet the demands of a school educational program and remain one of the most diverse and authentic scientific collections of temperate trees in the world.\nThe age of Kew only adds to the challenge: how does a curator not only maintain but hopefully improve upon a tree collection that has been tended for more than 250 years? The gardens at Kew date to 1731, when King George II's son, Frederick Prince of Wales, leased the estate and began to develop the grounds. After his death, his wife, Princess Augusta, continued his work, and in 1759, on the advice of Lord Bute, her horticultural advisor, she created a nine-acre botanic garden with the planting of several newly introduced trees that we now know as the 'Old Lions.' Some of these are still growing today, including a maidenhair tree (Ginkgo biloba) and a black locust (Robinia pseudoacacia). By 1768, the collection included almost five hundred hardy trees and shrubs, but it wasn't until 1840 that Kew Gardens was placed under direct government control and the first director, William Hooker, was appointed to restore and expand the arboretum.\nIt has been an amazing privilege to oversee such a collection, following in the footsteps of remarkable people like William Jackson Bean, the assistant curator of the arboretum between 1900 and 1922. He authored the monumental reference work Trees and Shrubs Hardy in the British Isles, which is now online (with regular revisions) as Trees and Shrubs Online, courtesy of the International Dendrology Society. Even within such a storied landscape, the collections are ever-changing. Managing those changes is the essential work of a curator.\nA landmark turning point for the arboretum occurred on the night of October 16, 1987, when a hurricane struck the southeast of England, wreaking havoc to trees and woodlands, felling over fifteen million trees in its wake. At Kew, over seven hundred mature trees were lost that night. I remember waking up to loud bangs and crashes and my steel dustbin rolling down the road. I got up to retrieve it and was concerned by the strength of the winds. The following morning, all came to light with the news showing images and footage of devastation across the south of England. \nI was a young supervisor in the arboretum at the time, and when I finally made the journey into work, I immediately went out into the landscape to see how all my arboreal friends had fared through the night. As I picked my way through the limbs and uprooted trees, all I could think was 'doom and gloom.' It took us over three years to finally clear away the fallen, damaged trees. As I look back now, I consider this hurricane to be one of the best things that happened in the twentieth century for trees in the United Kingdom. It raised public awareness of the importance of trees nationally. At Kew, a new plant exploration program was started to replenish the gaps in the collections created by the storm, and new arboricultural practices were developed to improve the health of the remaining trees.\nI was fortunate to be a part of the team sent to collect new documented seed material to rebuild the tree collections. The species on the target lists and the parts of the world that would be visited were determined by an audit of what was still represented in the collections after the storm, looking at the taxonomic and geographic weaknesses. The first expeditions were to western China, South Korea, Taiwan, the Russian Far East, and Japan, and the material brought back over the past thirty-four years has greatly enriched the diversity and provenance of the tree collections. Much of this has not been done alone. Working with colleagues at other arboreta around the world has been important for sharing ideas, collections, and stories.\nI have never been one for pushing the boundaries of hardiness, especially as we increasingly experience unpredicted weather patterns. Still, I have been able to plant and establish species that we could not have grown outdoors forty years ago: for instance, the Taiwan coffin tree (Taiwania cryptomerioides), Kashmir cypress (Cupressus cashmeriana), and the paran\u2021 and bunya pines (Araucaria angustifolia and A. bidwillii), both from the Southern Hemisphere. On my daily walks through the arboretum, I would look for locations to position these and others. As curators, we all have our favorite areas and genera of trees, but we must ensure that other parts of the collection aren't neglected. I found that the wire cages used to protect our young trees provided a helpful visual cue. The cages are retained for five years, so I would stand in the arboretum and turn 360 degrees. If I failed to see one of the cages, this would signal to me a target area for succession planting.\nSeveral new introductions into the arboretum come to mind as highlights. In the autumn of 1996, on a collecting trip to China, I was fortunate to be granted permission to visit Jinfushan, a mountainous preserve in the upper reaches of the Yangtze River, to see the Chinese silver fir (Cathaya argyrophylla). This species, discovered in 1955 by Chinese scientists, was something we had only heard about but never seen. We found it growing on the limestone bluff but could not collect seed, owing to a national embargo. Two years later, the embargo was lifted, and seed was distributed to forestry institutes and botanic gardens. The Forestry Commission's Bedgebury Pinetum was the first to grow this tree in the United Kingdom, and its curator gave me a two-year-old plant for our collection. This can be a miffy species and finding the best planting position can be difficult. More by luck than judgement, I got it right. The plant at Kew is now a beautiful specimen about twenty feet high. It has produced viable seeds, and the first generation of ex situ propagated seedlings has now been planted out in the arboretum, helping conserve this rare tree.\nAnother successful introduction is the Chinese hickory (Carya cathayensis). In 2008, on a trip to China to follow in the footsteps of Ernest Henry Wilson, I visited a market in Shanghai and saw nuts of the rare species being cooked and sold as candied pecans. We bought a kilo of uncooked seeds, and the propagator in Kew's nursery, after much experimental work, successfully germinated the seed and grew over twenty plants that are now sited in various locations across the arboretum. These are now gorgeous trees. They are very well-behaved, needing little if any formative training and producing a straight tapered trunk with an even distribution of lateral branches. The species is perfectly hardy in the United Kingdom.\nFor me, one of the main criteria for a successful and healthy treescape and collection is continual succession planting, maintaining a healthy population with generations of individual species, like a family, ranging from the great grandparents (the Old Lions) to the great-grandchildren (the newly planted trees this year). It was so rewarding to walk the collections seeing new introductions like the delicate Taiwan beech (Fagus hayatae), which we introduced as seed to the West in 1992, growing into strong, attractive specimens and enhancing the conservation value of the arboretum. Some of these, we hope, will be the Old Lions of tomorrow."},{"has_event_date":0,"type":"arnoldia","title":"Plant Rescue on the Cliffs of O'ahu","article_sequence":3,"start_page":11,"end_page":12,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25761","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170bb6d.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Sugii, Nellie","article_content":"In 2004, the last remnants of an exceedingly rare Hawaiian species, Cyanea grimesiana ssp. grimesiana, bloomed and set fruit in the wild. Known only from the leeward slopes of the southern Ko'olau Mountains on the island of O'ahu, this shrub is one of seventy-eight species within an endemic Hawaiian genus commonly known as hh. The species could be found surrounded by koa (Acacia koa) and other common forest trees, and it has been rare since it was first documented in the wild in 1819. Significant surveys occurred in the 1990s, and by 2004, only two mature wild plants remained, with no evidence of recruitment or any significant ex situ collections. The situation became dire.\nAt the time, I was several years into my career as a researcher for the Lyon Arboretum's Hawaiian Rare Plant Program, where I'm now the program manager. Our work focuses on rescuing and recovering Hawai'i's most critically endangered plants, storing germplasm for ex situ conservation, and providing plants for in situ restoration. Our micropropagation laboratory is central to this effort a surreal indoor space where more than 170 of Hawai'i's rarest and endangered plant species are grown collectively in tens of thousands of test tubes. I often describe it as 'plant conservation through the looking glass.'\nWhen the hh remnants flowered, our team worked with collaborators, including the Plant Extinction Prevention Program, the US Fish and Wildlife Service, and the US Army's Natural Resource Program, to plan for protecting the species in the micropropagation facility. Field biologists monitored the two plants. The flowers emerged as white, arching tubes, streaked with vibrant purple. The fruits then ripened into orange, fleshy capsules. The biologists carefully collected the fruit and brought it to the micropropagation lab for germination. We knew it was a heavy responsibility when the precious seeds arrived, but excitement ran through the lab as we sorted, cleaned, and prepped the seeds for in vitro seed sowing.\nMicropropagation gained recognition as a viable propagation method for commercial applications in the 1960s, but the technique was initially viewed suspiciously due to associated terms and applied technologies such as cloning, anexic seed sowing, ovulo culture, and organogenesis. To some, even at the Lyon Arboretum, these technologies seemed contrary to conservation theologies of preservation and genetic integrity. Yet micropropagation has gradually proven itself as a useful rescue and recovery tool. It can be used to germinate immature seeds and rescue embryos from aborted fruit. It's also used for cloning wild plants at risk of extirpation in order to preserve genetic representation and establish clonal lines of its seedling progeny for restoration.\nAfter the hh germinated in our lab, we learned that the final wild remnants had altogether succumbed the species no longer existed in the wild. This knowledge brought bittersweet feelings as we watched the seeds germinate in the petri dishes and eventually grow into seedlings that we placed into individual test tubes. We knew that it was now our responsibility to establish perpetuity for this species by establishing clonal lines of the seedlings through microcuttings and maintaining the in vitro germplasm collection until a safe and secure restoration site free of threats became available.\nApproximately 88 percent of the native plants on the Hawaiian Archipelago naturally occur nowhere else in the world. This rich biodiversity serves as a unique example of insular evolution, but its fragility is evident by the scale of species on the brink of extinction. According to listings by the US Fish and Wildlife, about one-half of the nation's threatened and endangered plant taxa are from Hawai'i. Of the five hundred Hawaiian species assessed for the International Union for Conservation of Nature's Red List, about 87 percent are classified as endangered or threatened. Let us not mention the hundreds of rapidly declining species that are missing from either list but are at risk of extinction.\nOn August 23, 2013, over nine years after the eventful collection date, I gathered at a site in the Ko'olau Mountains with a group of individuals involved in the conservation of Cyanea grimesiana ssp. grimesiana. A festive mood spread among us. We had long awaited the moment when we would bring this species and a few associated plants back to its native habitat, within the Mnoa Cliff Forest Restoration site. Our small group of friends and family even a few children made our way through a forest of an invasive bamboo that had taken hold in the area. A few of our team wore backpacks containing plants, and most everyone else carried trays of plants or tools in our hands. A space opened in the bamboo, and a pocket (or kipuka) of near-intact native forest appeared before us. For those seeing it for the first time, the beauty of the area took our breath away. We all acknowledged that the enclosure represented a new beginning for this hh.\nBy 2021, the original Mnoa Cliff plantings had matured. The hh plants flower and produce fruit, and the seeds are collected and sowed for restoration purposes or stored in our program's seed conservation laboratory. We have now stored thousands of seeds from the different plants, and we continue to maintain the original clonal lines in the micropropagation lab, with long-term cryopreservation being our future and final ex situ storage goal. With many hands and great effort, we have brought Cyanea grimesiana ssp. grimesiana back home."},{"has_event_date":0,"type":"arnoldia","title":"Thinking Outside the Quad","article_sequence":4,"start_page":13,"end_page":14,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25758","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170b36d.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Feldman, Carmia","article_content":"When Karyn Utsumi entered the University of California, Davis, majoring in environmental science and management in 2017, she didn't anticipate that she would eventually spend countless hours wearing waders and working with other students to restore a prominent water body on campus. Yet she knew that she wanted to turn her deep care for the environment into something that made a difference in her community. During her freshman year, she saw an announcement about the Waterway Stewardship internship with the UC Davis Arboretum and Public Garden. She applied and was thrilled to be selected.\nThe UC Davis Arboretum and Public Garden spans the entire 5,300-acre university campus, with a historic arboretum, founded in 1936, at the center. By applying the management and engagement principles of a public garden to the campus at large, the university aims to enhance how the entire Davis community views and interacts with its environment. Our student internship program, which Karyn joined, is our top initiative to do just that by developing the next generation of environmental leaders. The program is called Learning by Leading\u00aa. Students gain leadership and technical skills as they tackle critical environmental issues with real-world, hands-on projects. As students progress through the program, they take on more responsibility through our mentor-supported 'leadership ladder.' Students start as learners and then can work through a succession of leadership positions, including project leader, team leader, and apprentice.\nFor students in the Waterway Stewardship internship, their living laboratory is the Arboretum Waterway, a creek-like body of water that runs through the historic section of the arboretum. The waterway is part of the campus stormwater drainage system and is dammed at both ends. While it resembles a creek, the Arboretum Waterway is effectively a pond, which means that it comes with common pond issues: nutrient-rich water and unsightly algae formation. After Karyn was hired as her team's coleader during her junior year, she led her interns in developing a floating wetland with sedges and other native plants that take up nutrients from the water as they grow. From afar, the planting resembles a green island. She worked hard to create consequential experiences for her team, learning to see and celebrate each member's unique skills.\nOver seven hundred students have now gone through the Learning by Leading program since it began in 2008. Another student, Ricardo Black, transferred to Davis from Los Medanos College, a community college in Pittsburg, California, for his junior year in the fall of 2019. He became a student leader for our Habitat Horticulture team, which enhances the suitability of campus gardens for native pollinators and other wildlife. Ricardo and his team worked in the Pollinator GATEway Gardens in the arboretum proper. A series of GATEway Gardens have been designed collaboratively with academic departments to showcase their research and teaching to visitors. The Pollinator GATEway Gardens, highlighting plants important for native bees, butterflies, hummingbirds, and other pollinators, were created with the nearby School of Veterinary Medicine. The project aligns with the school's research on the interconnections between the health of people, animals (both domestic and wild), and their environment.\nRicardo's leadership skills were tested when the pandemic forced our normally hands-on, outdoor internships into a virtual format. He demonstrated fast, adaptive leadership as he navigated his team through the initial unpredictable months of the pandemic. He found that it became even more essential to develop peer-mentor relationships, which encouraged his growth as a communicator. Ricardo says, 'During the program, I was put in a position where leadership and innovation skills were needed to make things work in an environment that was unpredictable and always changing due to the pandemic.' Similarly, Karyn credits the Learning by Leading program for shaping her into the collaborative leader she is today. When she started the internship, she told herself, 'I need to work hard and figure everything out by myself.' Then, as she progressed through the program and gained leadership experience, she realized that strength comes through working together.\nKaryn also says that Learning by Leading helped her discover her twin passions for restoration and environmental education. She graduated in the spring of 2021 and immediately was hired by two local environmental organizations: the Putah Creek Council, where she organizes community volunteers to do creek restoration work, and the Solano Resource Conversation District, where she serves as an environmental educator. Karyn's growth as a leader and her impactful postgraduate jobs exemplify the power of reimagining the traditional university campus. All university campuses are more than lawns, sidewalks, and buildings they can be spaces where tomorrow's environmental change-makers learn to lead."},{"has_event_date":0,"type":"arnoldia","title":"Water Comes First","article_sequence":5,"start_page":15,"end_page":17,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25757","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170af6a.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Bartlett Jr., Robert A.","article_content":"My decision to transform the R.A. Bartlett Research Laboratories and Arboretum into the living museum that exists today was centered on the need for water. My father, Robert Bartlett Sr., purchased the property in 1965, a few years after he became president of the family business, Bartlett Tree Experts. He intended for the 350-acre property, nestled in the rolling hills outside of Charlotte, North Carolina, to serve as a research laboratory and training center for the growing company.\nTo that end, the company established facilities and plots where staff scientists conducted experiments on plant care and pathology. Previously, this work had been performed in Stamford, Connecticut, where my grandfather had set up our first tree research laboratory and training facility in 1913. A portion of the original site still exists today as the Bartlett Arboretum and Gardens, although it has no affiliation with the company.\nPreviously, the Charlotte property had been a working horse farm with large fields and pastureland. When my father acquired the land, he planted azaleas (Rhododendron) and hollies (Ilex), along with other plants that form the basis of the collections we have today. In those early years, I remember seeing young trees begin to establish themselves and rise above the forage grasses.\nThe climate near Charlotte allowed the cultivation of species common in both northern and southern gardens, which was important since we had field offices throughout the United States (and now Canada, the United Kingdom, and Ireland). Still, the summers in Charlotte are hot and humid. To maintain the collections, we pumped water from one of three existing ponds to provide irrigation, but at first, the capabilities were technologically limited. The earliest systems consisted of gasoline-powered pumps sitting on the shore of the ponds and serving manually operated spigots. Remnants of those systems can still be found on the property, and portions of their piping are still in use today.\nDuring the first thirty years in our Charlotte location, it was apparent that some of the plants were beginning to suffer from our limited irrigation capabilities. In particular, the collection of azaleas that my father had started planting on an eighty-foot hill, now affectionately called Rhodie Hill, required extensive watering. In midsummer, it was challenging to stay ahead of the heat, especially because the water had to be manually hauled up the paths that wind around the hill.\nThe impact of a changing climate also became more apparent at this time. When my father bought the land, the property was categorized by the US Department of Agriculture as being in plant hardiness zone 7 (meaning the average minimum temperatures fell between 0 and 10\u00a1F); however, it is now considered zone 8 (averaging between 10 and 20\u00a1F). Due to changes in the weather patterns, longer dry periods developed, and more dramatic swings in rainfall became the new normal. It was clear that we could no longer sustain our collections without investing in a state-of-the-art irrigation system.\nAfter my father passed away in 1998, we began to make a significant investment to help maintain and develop the property. It would continue to serve as a research station and laboratory, complete with a training facility for clients and arborists and a diagnostic clinic where our researchers process thousands of plant and soil samples sent by our field offices. At the same time, we were determined to continue building the collections into a world-class arboretum. With this goal in mind, we decided to put in an irrigation system that could provide consistent water to the growing collections.\nWe installed a new distribution system to feed the early network of pipes and facilitate manual watering capability in adjoining areas. Most importantly, the system directed a large volume of water to one of our ponds. Now, with the ability to keep a single, large reservoir of water full at all times, the Research Lab and Arboretum was primed for much more extensive, and automated, irrigation operations. In 1999, we began installation of the first automated system. It allowed us to direct a precise amount of water overnight to specific areas on the property. The collections grew like never before. The system also made new locations available for dedicated research plots. Automatic irrigation was a game changer.\nAt that point, we began to strategically build our collections. We launched collaborations with other arboreta and research institutions across the globe and started adding to the diversity of our cultivated plants. Today, the collections are expansive, consisting of over twenty-six thousand accessioned plants in fourteen major groups. We have one of the largest collections of holly in the United States, along with extensive collections of elm (Ulmus), crape myrtle (Lagerstroemia), maple (Acer), witch-hazel (Hamamelis), linden (Tilia), and boxwood (Buxus). Seven collections are accredited through the Plant Collections Network, including the largest collection of Magnolia cultivars in the world. \nAmong the collections, those which were established early and added on to over the years continue to be among the most satisfying for me to watch through the year. Our main grouping of magnolias borders Youngblood Road, a two-lane highway that passes the arboretum. When you drive around the corner and see the magnolias in bloom, the sight of the different colors almost takes your breath away. There is just about every shade and hue of purple, pink, white, and yellow that you can imagine. Rhodie Hill is another favorite. The hill comes alive in a kaleidoscope of spring color, and with mature specimen trees overhead, the winding paths offer beautiful surprises around every corner.\nWe have now begun focusing on wild-collected plant material, especially prioritizing species of conservation concern. One of the plants that we are playing a role in conserving is a rare North American species known as the pyramid magnolia (Magnolia fraseri var. pyramidata). In an effort to understand the distribution of this species and increase documented holdings in cultivation, our arboretum has partnered with The Morton Arboretum, the University of Florida North Research and Education Center, the Chicago Botanic Garden, the Atlanta Botanical Garden, and the US National Arboretum to scout populations, assess their health, and collect seed (when present) for propagation and distribution. Through collaborative efforts like this, and with other strong networking partners such as the Arnold Arboretum, Longwood Gardens, and many others, we have made conservation of rare species a new part of our mission. \nLooking at all the natural beauty established here, visitors may find it easy to forget that this is a relatively young arboretum. We pride ourselves on the ability to adapt with the times and use our natural water resources to maintain the vitality and health of our collections. The key and catalyst to our success has been access to water and having the irrigation needed to help the plants thrive. Without it, we could not have created this botanical wonderland in such a short amount of time."},{"has_event_date":0,"type":"arnoldia","title":"Metasequoia glypotostroboides","article_sequence":6,"start_page":18,"end_page":19,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25762","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d1708125.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Jahren, Hope","article_content":"If you head north, far above the Arctic Circle, you will find yourself in a land of blue sky, white snow, and gray ice. There will be pockets, here and there a lonely island, a sloping valley that are dry, dusty, and desolate. Dig down, through a crust of lichens, take out the smooth stones underneath, and burrow into the shaley, ancient mud. When you get to layers no less than forty million years old, you will find conifer needles. Not only that, you will find twigs, branches, cones, and even whole trunks, dusted in ancient sap. I have seen this myself, during the odd, dream-like hours that are born of twenty-four-hour light.\nForty-five million years ago, at 79\u00a1 north latitude, an immense conifer forest stretched in every direction, across what is now Canada, Alaska, and Siberia, quite close to today's North Pole. The idea of a forest so far north is nothing short of fantastic: today, the tallest plant in the region is a pussy willow and a stunted specimen at that. The temperature and the rainfall above the Arctic Circle were certainly very different forty million years ago. Still, one thing has not changed: total light for three months, soon followed by three months of total darkness. No modern trees can tolerate these conditions, yet forests once thrived under this ridiculous annual regime. Foremost among the trees was Metasequoia. We recognize them from their needles fossilized but so loose that they fall through your fingers like confetti.\nUntil 1948, most scientists assumed that Metasequoia was extinct, based on fossils from lower latitudes. That was the year the Arnold Arboretum received a package from Hu Xiansu, who trained at the Arboretum and returned to China with his doctorate in 1925. Hu sent bushels of seeds and other botanical materials, and he documented that they had come from wait for it live Metasequoia glyptostroboides growing in central China! Some of these seeds became the full-grown, magnificent 'dawn redwoods' that now stand throughout the Arboretum (accessions 3-48 and 524-48).\nBecause of these seeds and the trees they became, I knew something about the fossils that we excavated in Canada that I would never have known otherwise: ancient Metasequoia trees were deciduous. Deciduousness is a special type of dormancy meant to decrease the stress of maintaining leaves through the winter. This trait, uncommon in conifers, would make all the difference as the trees prepared for the extended Arctic darkness."},{"has_event_date":0,"type":"arnoldia","title":"The Third Fifty Years of the Arnold Arboretum","article_sequence":7,"start_page":20,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25756","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170af26.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Round the bend on Hemlock Hill Road and look across Bussey Brook and Kent Field to the north. Your eyes will skim a patchwork of conifer textures, colors, and forms. Among the trees is an upright individual with a rather abrupt taper at the top, the Arnold Arboretum's largest giant sequoia (Sequoiadendron giganteum, accession 1320-72*A), now standing eighty-three feet tall. After crossing the brook and walking up the slope, you'll see that the wide bole (almost five feet in diameter) begs to be hugged. Shift your gaze up along the orange bark to the sky, and you'll see the tree's candelabra-like branching pattern. Most of the branches seem normal, erupting out of the main stem at right angles,\nbut if you step back and keep your eye on the crown, you'll see an odd conglomeration where one branch over another attempted to bend skyward.\nBack in 1948 (the same year that Metasequoia glyptostroboides, the dawn redwood, arrived in North America), a supporter of the Arboretum, Chandler Hovey, collected giant sequoia seedlings from California and planted several near his home in Brookline, Massachusetts, a stone's throw from the Boston College campus. In 1972, in honor of the Arboretum's centennial, Boston College which had recently acquired the Hovey property donated the tree. That spring, a twenty-four-year-old, forty-two-foot-tall, pointy-topped tree was dug, transported, and transplanted in its current spot in the conifer collection. The magnificent specimen survived, but its central leader died due to transplant shock, leaving an oval-shaped form for decades. A new leader eventually took over: a dog-legged branch that formed fifty years ago from the initial crown's tip, some forty-three feet above the ground. I'm certain that the wooden rings within that branch would reveal not just what was going on with that single tree but the surrounding Arboretum landscape as well.\nThe First Fifty Years\nJust as the sequoia's history is written within its rings, branches, and form, the Arboretum's landscape and collections reflect a history rich in dramatic events and subtle ripples. Much has been written about them, and Charles Sprague Sargent's 'The First Fifty Years of the Arnold Arboretum' describes the first five decades with aplomb. At the time of Sargent's writing in 1922, Harvard's tree museum (founded in 1872) had expanded from 125 to 250 acres. Frederick Law Olmsted had reimagined Benjamin Bussey's farm with carriageways and pathways, collection areas and viewsheds. Sargent and his team transformed the landscape into a composite of taxonomic tree groups and research plantings, including an intensely cultivated shrub and vine collection, all nestled among a few natural and naturalized woodlands.\nBy 1922, botanical exploration particularly of East Asia and North America and horticultural exchange yielded a living collection of over five thousand taxa growing at the Arboretum. The institution was well on its way to meeting its initial charge (a nascent collections policy, if you will) to cultivate every tree, shrub, and vine hardy in Boston. While many of the plants were botanical taxa, including wild-origin species newly cultivated in North America, there was no shortage of infraspecific forms and varieties that we would now call cultivars. The herbarium of two hundred thousand sheets complemented a thirty-five-thousand-volume library and archival collection of nearly ten thousand photographs. With these integrated living, preserved, and archival collections, the Arnold Arboretum had become an international destination for scholars of woody plants.\nYet Harvard's tree museum was not just for the botanical connoisseur. This gem in Boston's Emerald Necklace of parks provided open space to an expanding and diversifying city. Because of the 1882 arrangement where ownership of the land shifted from Harvard to the City of Boston (who then leased the property back to the university for at least one thousand years), the space would be secured in perpetuity as both a scientific enterprise and public open space, free for all to enjoy. Without this arrangement some 140 years ago, I doubt if the Arnold Arboretum would exist today, or if it did, if we would recognize it in its current form. Had it remained a nonpublic, university-owned research station, I can imagine acres by the dozen being peeled away and sold with each-and-every economic crisis. If purely a municipal park, even if well maintained, it would not house one of Earth's most notable collections of woody plants. Luckily, these are just what-if scenarios.\nPerhaps knowing his grip upon the Arboretum's helm would not last much longer (though it did, for another four years), Sargent ended his half-century assessment with a few bold charges for his successors. Global environmental change was apparent to him, particularly the challenges to trees and forests worldwide. Thus, Sargent called for continued and ambitious documentation of forests in Asia and the tropics, as well as rigorous scholarship in forest pathology, entomology, and genetics. Within the Arboretum landscape, Sargent felt that a rose and a rock garden would be essential additions, no doubt to provide space for new collections development while simultaneously enhancing the horticultural display. Despite the growth of the initial Arboretum endowment from $103,847 to $808,175, Sargent knew that additional resources would be required not just for these new initiatives but to maintain current operations. Thus, he curtly ended his fifty-year report with one sentence: 'Only a larger endowment is needed to make possible these Arboretum activities and extensions.' Following his death in 1927, the Sargent Memorial Fund would raise over a million dollars.\nThe Second Fifty Years \nThe Arboretum's second half-century was dramatic and dynamic. The institution weathered a global economic depression, multiple leadership changes (one supervisor and three directors), the catastrophic hurricane of 1938, as well as another World War. There was also the Controversy (as it was referred to), which amalgamated the university's herbarium and botanical library collections (including most of the Arboretum's) under one roof in Cambridge. This coincided with the cessation of the Bussey Institution, which had opened as Harvard's center for horticultural and agricultural education in 1871, on property adjacent to the Arboretum. The institute grew into a center for genetic and cellular research. By the 1930s, most of the on-site scholarship in the Arboretum's living collections had waned. Likewise, the Arboretum's fieldwork in temperate areas, particularly to acquire germplasm to grow in the living collections, ceased almost entirely. However, botanical exchange of seeds persisted, with the 1948 acquisition of the Chinese dawn redwood, Metasequoia glyptostroboides, being one of the most celebrated feats even to this day.\nThe discipline of horticulture often considered the art and the science of growing plants matured in the mid-twentieth century. The genetic improvement of ornamentals hit a stride, as did advanced ways to propagate them clonally. As a result, cultivars (first given formal recognition in 1952) wantonly tumbled out of nursery catalogues and into gardens, parks, and other managed landscapes. The Arboretum's living collection was no exception.\nIn 1970, Donald Wyman, horticulturist in charge of the Arboretum from 1935 to 1970, wrote in these pages about the Arboretum's goal to improve the curation and care of the collections, and to use the collections as a living laboratory for horticultural introductions. Species plucked from the wilds in the Arboretum's first half-century would be assessed for their garden worthiness in the second. Novel hybrids, like crabapples (Malus) and forsythia (Forsythia) created by Karl Sax (a professor who then served as director from 1947 to 1954), were given growing space, with many introduced as cultivars after evaluation. Arboretum selections, and those from sister institutions and industry, were grown side-by-side, their performance recorded in Arnoldia and other publications.\nJust like there were changes to what the Arboretum grew in the collections and why, there were changes to where they were grown. The 1942 acquisition of the Case Estates in Weston, Massachusetts, provided a welcome relief valve for the space-cramped Boston collection. Shortly thereafter, several ornamental collections in Boston underwent redesigns: Landscape architect Beatrix Farrand's Azalea Border along Meadow Road added dramatic color and space for the deciduous Rhododendron that were performing poorly elsewhere. Crabapples, the dandy of mid- to late-twentieth-century landscapes, replaced most of the hawthorns (Crataegus) on Peters Hill during renovations from 1948 to 1952. The construction of the Dana Greenhouses in 1962 provided a sophisticated station where propagator Alfred Fordham could conduct his many experiments and publish them widely.\nCollections of the Third Fifty Years\nThe centennial in 1972 arrived with fanfare and excitement. Dick Howard, director since 1954, began his 1971 annual report to the Harvard University Provost by underscoring the Arnold Arboretum's essential service role to the City of Boston, particularly to local communities. Maintaining the Arboretum required considerable resources that were worth the expense and investment, and caring for the collections was his 'priority responsibility.' Thus, irrigation projects in both Boston and Weston would alleviate some of the growing and unmet demands for water. A bucket truck was added to the fleet, which made pruning or removing old, senescing 'stag-headed' trees easier. To replace some of the removals, horticulturists planted out nearly nine hundred specimens, completing a cycle of rejuvenation and renewal. Anticipating future databasing, Howard noted that the plant records office had wrapped up a major inventory campaign to assess and field-check every specimen in the collection.\nOver the Arboretum's third fifty years, the institution would be led by four directors: Richard Howard's tenure ended in 1978; Peter Ashton led from 1978 to 1987; Robert Cook from 1989 to 2009; and William (Ned) Friedman became director in 2011. During this time, the Arboretum experienced dramatic changes, as did the living collections. Staff actively contemplated what to cultivate, where to grow it, and how to do it better.\nMajor anniversaries like a centennial can elicit reflections and ambitions, so it is no surprise that shortly after Peter Ashton became the director in 1978, strategic planning was underway. One broad initiative, a restoration plan, included a substantial section for what should be in the collections. A formal living collections policy the first for this Arboretum and most botanic gardens was also published in 1979, remaining in force for almost thirty years. In this latter document, the Arboretum established and codified ambitious goals: to acquire all known woody species hardy in Boston (no different from the original charge of 1872); to have three individuals of each species; to prioritize wild-provenance plants above those of garden or nursery origin; and (assuming they met specific requirements) to continue to maintain taxa at infraspecific ranks (including cultivars, although these were considered lowest in any hierarchy).\nTo complete the collections, the 1979 restoration plan outlined the addition of over 2,900 taxa, spanning 90 families and 363 genera. These desiderata came almost exclusively from identifying which plants in the 1940 edition of Alfred Rehder's Manual of Cultivated Trees and Shrubs Hardy in North America were missing from the collection. To launch the initiative, staff set an ambitious goal of acquiring 1,500 taxa in the first five years.\nEven before strategic planning of what to add, the Arboretum reconsidered where new material would come from and how to acquire it: collectors would return to the field. In 1977, the Arboretum embarked on its first major collecting trip in some forty years, sending taxonomists Stephen Spongberg and Richard Weaver to South Korea and Japan for six weeks. In 1980, following the heels of the restoration plan, Weaver botanized in the Soviet Union, while Spongberg participated in the three-month-long Sino-American Botanical Expedition, which involved a team of thirteen Chinese and American collaborators. (China had not been visited by Western botanists since before the revolution in 1949.) The era of fieldwork had returned.\nThrough the 1980s and early 1990s, the infusion of wild-collected material from some seventeen expeditions occurred at a scale not seen in fifty years. In some years, the Arboretum sponsored multiple collecting trips. Destinations included those known to yield hardy material such as northwestern Hubei Province, China, and the Appalachian Mountains of the American Southeast, as well as fringe regions like North Africa, Mexico, and Taiwan. Coincident with fieldwork, the Arboretum also received new material from sister institutions, often selected from their annual seed lists (known as index semina).\nAlthough the restoration plan advised against 'returning to what must have been almost a jungle by the end of Sargent's tenure as Director,' there was no discussion as to where some five thousand new plants (an increase by approximately 30 percent) would be sited in the collections. Thus, limitations in capacity and resources facilities, staffing, and space hindered the restoration's full success. For one, the Arboretum lacked the facilities to propagate and produce the sheer magnitude of material arriving in such a short period. The plant records database is replete with notations from index cards of whole flats of accessions that perished due to the lack of production space (many were placed in the shade below the benches). Gary Koller, Wyman's successor as the lead horticulturist, has told me how, due to severe space constraints in the collection in the 1980s, sibling plants of the same accession were planted together in tight triads, about five feet apart. Only a few of the triads remain today, primarily sited along the roads and perimeter of Bussey Hill. Deaccessioning plants was taboo, so there were few other alternatives. The 1979 restoration document was successful as an acquisitions plan yet perhaps too ambitious given practical considerations.\nA decade later, following changes in Arboretum leadership (Robert Cook became director in 1989), a Living Collections Long-Range Planning Committee returned to the process of thinking about the collections. In 1991, the committee completed a planning document, edited by Stephen Spongberg, which acknowledged that the 1979 restoration may have been na\u2022ve. The committee noted the challenges in adopting a comprehensive collection (meaning one of every taxon) versus a synoptic or broadly representative collection. They observed that it would be difficult to preserve the integrity of the Arboretum's historic landscape in light of the aggressive drive to acquire new material. Nevertheless, the plan ended with a reaffirmation of the same ambitious collection policy goals articulated in 1979.\nTo accommodate this expansion while remaining sensitive to the Olmsted design (by not transforming the collections into a dense forestry plantation), the 1991 plan called for the prudent review and deaccessioning of low-value and out-of-sequence material. The authors proposed a long-term review process that would finally deal with many of the growing pains that had affected the Arboretum since (and perhaps prior to) the death of Sargent. Although there were no estimates of how many plants could be deaccessioned, the authors stated that such subtractions would be insufficient to accommodate the necessary expansion. The 1991 plan estimated that all Arboretum property must be designated for the purpose of housing an expanded collection, including the entirety of Peters Hill, Bussey Brook Meadow (formerly called the South Street Tract or Stony Brook Marsh), Weld Hill (formerly Weld-Walter Street Tract), and the Case Estates. Space was not the only resource required: the plan identified new staff positions necessary for curation, horticulture, and the greenhouse and nursery.\nShortly after the 1991 plan was completed, it was put on hold following a reorganization of the Arboretum's administrative structure in early 1992. A new Living Collections Department was created, with Peter Del Tredici leading. The ambitious goal of the 1979 and 1991 plans to form a comprehensive collection was admittedly unrealistic and abandoned. Instead, as Del Tredici outlined in 1994, collections development would take a more focused or prioritized approach. During the early 1990s through the mid-2000s, special recognition was reserved for conservation-status species (particularly those maintained in collaboration with the Center for Plant Conservation). As a theme, the floras of eastern Asia and eastern North America were given priority, particularly genera like Acer (maples) and Fagus (beeches), which became two of the initial five collections nationally accredited through the Plant Collections Network. (The Arboretum now has eight accredited collections.) The recently established North America China Plant Exploration Consortium (NACPEC) became a pipeline for novel germplasm from China. From 1991 to 2006, the Arboretum mounted six expeditions to China, two under the NACPEC flag, including the 1994 expedition to Hubei that infused the collections with new material like the paperbark maple (Acer griseum), which had most recently been collected by Ernest Henry Wilson in 1907.\nWithin a year of joining the staff in 2007, I organized a team to update collections goals and codify them in a new living collections policy. The scope of the collections would remain synoptic, with the highest priority assigned to core collections, such as the nationally accredited collections and conservation-status holdings. Historic lineages would be maintained through repropagation, while targeted acquisitions of cultivars would meet trialing, display, and research needs. The new policy (and its subtle revisions over the past fifteen years) prompted the review and subsequent deaccessioning of excessive or low-value accessions, as well as the repropagation of valuable lineages that had gone unnoticed.\nFieldwork continued, with another six expeditions occurring between 2007 and 2015, including a NACPEC expedition to the Qinling Mountains of China in 2010 and a more focused collecting of live oak (Quercus virginiana) from the northeastern edge of its range in Virginia in 2012. In 2015, the Arboretum launched the Campaign for the Living Collections, an initiative that followed several years of planning from the Living Collections Advisory Board. The campaign articulated a list of nearly four hundred target taxa, each linked to one or more priority themes found in the collections policy. Since the campaign launched, some twenty expeditions to destinations in the United States, China, Japan, and the country of Georgia have yielded over half of the desiderata. The COVID-19 pandemic paused expeditionary work for 2020 and 2021.\nWhile the Arboretum embarked on exactly fifty named expeditions over the past fifty years, plants of cultivated origin were added to the collections (or maintained) for their invaluable ornamental characteristics, stress tolerance, and other novel traits valued in managed landscapes. Cultivars of trees continued to grow alongside their wild-origin brethren particularly in the Rosaceous orchards of Peters Hill while new shrub cultivars appeared in the Bradley Rosaceous Collection (dedicated in 1985), the Leventritt Shrub and Vine Garden (dedicated in 2002), and other landscapes. In 1972, 14,058 plants grew in the Arboretum's collections in Boston, and only 14 percent were of wild origin. As of this writing, 44 percent of the 15,939 plants in the collections were derived from wild populations, and if one excludes over 2,700 accessioned plants in the natural areas (such as Hemlock Hill, which is a mix of wild and planted hemlocks), 53 percent of the collections are from the wild. That is quite the illustration of focused and deliberate collections development.\nDesigning the Collections\nPerhaps the most significant outcome from the 1979 restoration plan was the recognition of historical planting areas, as articulated loosely using the Bentham and Hooker linear sequence. Richard Weaver created maps for each family and major genus, using red colored pencil to illustrate where new plantings should go (or, in some cases, errant shrubs should be returned). This reordering was meant to fix what were perceived as random horticultural plantings, particularly those from the mid-twentieth century.\nAll gardens need redefinition from time to time, and many areas within the Arboretum received edits over the past fifty years. For instance, Rhodie Dell the collection of broadleaved Rhododendron along Bussey Brook at the base of Hemlock Hill was renovated in 1990 with the Davison Path laid out by Julie Moir Messervy. The landscape around the Hunnewell Visitor Center received a new look by Carol Johnson after the building was renovated in 1993. In 2007, Beatrix Farrand's Azalea Border along Meadow Road received an infusion of new material following the removal of declining individuals.\nOne of the major goals Sargent described in 1922 was the creation of a rose garden, and in 1985 the Arboretum made good on this promise. A gift by Eleanor Cabot Bradley and an innovative design by Gary Koller created the Bradley Rosaceous Collection. Located near the ponds and replacing the existing shrub collection (where many of the Rosaceous shrubs grew already), this semi-formal garden adjacent the Forest Hills Gate became and continues to be a public gathering space and programming site. Updates completed in 2011 (by Julie Moir Messervy) improved circulation and display potential, and two wrought-iron arbors designed by Peter Andruchow added spaces for climbing roses.\nWhile the Bradley created a significant destination for visitors, the diaspora of shrubs and vines from the earlier shrub garden led to a problem. Many of the vines were moved to chain-link fences on the perimeter, becoming challenges to maintain, while sun-requiring shrubs now grown in the shade under their arboreal cousins did not always fare well. To ameliorate this dilemma, the Arboretum needed a new shrub and vine collection, and with a gift from Frances Leventritt, the Victor M. and Frances Leventritt Shrub and Vine Garden was created in 2002. Designed by Reed Hilderbrand, this formal garden would house sun-loving shrubs and vines on property to the north of the Dana Greenhouses, on space previously occupied by the old hedge and dwarf conifer collection. Unlike other areas of the Arboretum's collections, the shrubs and vines grown here were to receive intense horticultural care and inspire ideas for home landscapes.\nPeters Hill, often neglected due to a lack of resources and its distance from the hub of operations, began to receive attention starting with a curatorial review in 1993. Low-value plants were deaccessioned; new plantings (particularly crabapples and deciduous gymnosperms) followed; and a bus turnaround at the summit was removed and renovated to support plant collections in 1997. Another major change occurred in 1996 when the South Street Tract was combined with land owned by the Massachusetts Bay Transportation Authority and the City of Boston, creating what is now known as the Bussey Brook Meadow, a twenty-six-acre urban wild with the Blackwell Path connecting Forest Hills Station to the South Street Gate.\nStewarding the Collections\nIn his 1971 report, Richard Howard noted that his highest priority was the maintenance of the living collections. His successors possessed the same agenda, mustering resources to support them as creativity and windfall allowed. Over time, work at the Case Estates waned to the point that by 1991 the horticultural staff in Weston shifted permanently to care for the collections in Boston. (The final sale of the Case Estates occurred in 2017.) Ongoing growth in the Arboretum's endowment, particularly during the capital campaign ending in 2000, allowed further staffing increases, and restricted endowments for areas like the Bradley Rosaceous Collection and the Leventritt Shrub and Vine Garden funded exclusive and dedicated horticulturists to care for each high-maintenance area.\nAnother major shift in resourcing occurred with the launch of the Landscape Management Plan in 2008, a charge led by Richard Schulhof (the deputy director) and implemented by Stephen Schneider (then the manager of horticulture). Recognizing the value of having already designated horticulturists in several areas, such as the Shrub and Vine Garden, the Landscape Management Plan expanded the perspective to all areas of the Arboretum landscape. The landscape was divided into zones, with individual horticulturists assigned to steward each according to goals specific to each area. The plan also directed the work of arborists as they rotated through the collections, and landscape staff as they maintained meadows, turf, and pathways.\nIn addition to performing the day-to-day care of the collections, horticulturists must contend with periodic natural disasters, pests, and diseases. Though not as cataclysmic as that infamous and unnamed hurricane that struck in 1938 (which destroyed some fifteen hundred trees), the 1997 April Fool's Day storm dumped over two and a half feet of snow on a collection previously plagued by past droughts. Over four hundred trees had to be removed that season, while another thirteen hundred remained but required arboricultural care. Pathogens and pests are a persistent threat to the collections. For instance, in the 1980s and 1990s, phytoplasmas plagued the lilac (Syringa) collection, and in 1997, hemlock woolly adelgid arrived at the Arboretum's doorstep to forever change the face of Hemlock Hill, a unique natural landscape where black birch (Betula lenta) are slowly replacing the hemlocks (Tsuga). And, in 2018, many old beeches (Fagus) were removed due to decline caused by the arrival of beech bark disease. All three of these collections the lilacs, hemlocks, and beeches are nationally accredited, so their stewardship in response to these outbreaks is especially significant. The Landscape Management Plan includes response plans for disaster and plant healthcare issues like these.\nIn late 2019, Andrew Gapinski, as manager of horticulture, transformed the third edition of the Landscape Management Plan into a dynamic, digital format known as the Landscape Management System. As part of the system, a smartphone and desktop application called ArbManager replaced the paper forms (the 'green cards') exchanged between horticultural and curatorial staff to communicate about work requests, while an internal website, ArbDashboard, synthesized horticultural and plant records data into a map-based system. Both of these tools provide living collections staff instant access to collections-care directives, whether they are in an office or fifty feet up a tree and accessing the information from a phone.\nRecording the Collections\nThe Arboretum is replete with uniquely accessioned plants, each richly documented with source histories, observations, photographs, herbarium specimens, and maps a tradition dating back to the institution's founding. In Howard's 1972 annual report, he noted how the card catalog entries the original paper database for the living collections, if you will were incorporated into the Plant Records Center of the American Horticultural Society. This initial digitization effort was championed by Howard when he was president of the American Association of Botanical Gardens and Arboreta. The shared database gave gardens the chance to store computerized records off-site (as a preservation initiative) and recall specialized lists of plants on demand (for instance, all plants in a given location within a garden).\nIn 1985, the Arboretum's plant records and systems (including definitions, workflows, and philosophies) seeded a new database eventually called BG-BASE. The Arboretum now had local access to its data, which revolutionized how the Arboretum and finally other gardens curated their collections. At first, the database only included living plants; however, funding from the Institute of Museum and Library Services (IMLS) in 2010 allowed staff to integrate legacy data from old index cards into BG-BASE, providing access to historic collections that had long ago perished. An earlier IMLS grant, in 2001, enabled the digitization of records for some fifty thousand vouchers from the herbarium of cultivated plants, adding even more data and research value to plants that grow or grew in the living collections.\nHand-drawn maps had recorded the locations of plants growing in the collections since 1938, and in 1987, cartography went digital due to support from IMLS. Initially, AutoCAD served as the digital platform; however, in 2010, the platform shifted into ESRI ArcGIS, a more robust geographic information system. This change was timely, as the IMLS grant in 2010 also allowed for the scanning and georeferencing of some two thousand hand-drawn maps, providing staff the ability to view like a digital flip-book the historic collections over time. Coincident with the legacy of mapping has been the annual inventory process, whereby all accessioned plants are field-checked on a five-year cycle. Whereas earlier field observations required paper cards, notebooks, and copies of maps, the current team led by Kyle Port, the manager of plant records, employs live connections to the database in the field using laptops and tablet computers. I wonder what Professor Sargent would think if he could witness such activities in action!\nWhile countless other initiatives over the past fifty years led to curatorial reviews and data acquisition, one final, and significant, venture was a multi-year verification project funded by the National Science Foundation in 1984. This project led to the vouchering (using herbarium specimens) of the living collections. The vouchers were then distributed to taxonomic specialists around the world who verified the identity of each plant. The effort yielded positive (as well as negative) identifications and fostered international research interest in the living collections.\nUsing the Collections\nWhile this article mostly reflects the living collections and their change over time, to leave out access and use would be a grave mistake. The Arboretum is not a private collection but is, in fact, very public. Due to the porous nature of the Arboretum, visitor counts have always been a rough guess. Until recently, estimates of annual visitors were in the 'hundreds of thousands,' which at the time may have been accurate. However, a people counter at the popular Arborway Gate one of more than a dozen entrances tallied some 825,000 hits from September 2020 to September 2021. This number includes ins-and-outs as well as pass-throughs, yet even with a conservative estimate of half this total (equal in and out hits) of 400,000 visitors at this single gate, it is safe to assume that well over a million people, and perhaps over twice that number, visit the Arboretum each year.\nAfter the Bussey Institution ceased to exist and much of the herbarium and library migrated to Cambridge, it had become more difficult for the living collections to readily serve scholars. However, engaging scholars to use the living collections has long been an area of interest of mine, even before joining the staff, and was one of the reasons I was hired into this role fifteen years ago. Luckily, much work had been initiated before my arrival. Five decades of field exploration yielded a collection rich in botanical diversity and wild provenance: research specimens little different from what a scholar could find in the natural environment. Year-over-year improvements in horticultural care provided healthier plants available for study. Ongoing vouchering, verification, and inventory initiatives add rich documentation to the plant records, all of which have been searchable online for over two decades.\nFunding also helped bring scholars to the collections. In 1988, initial support (and later an endowment) provided by George and Nancy Putnam created the Putnam Fellowship specifically for those conducting independent research and project work using the collections. These and other competitive awards have helped to remove financial barriers that might otherwise prevent research from occurring.\nLastly, in 2011, the Weld Hill research facility opened. While it was initiated and built during the administration of Bob Cook and opened and staffed shortly after Ned Friedman became director, the facility was inspired by Peter Ashton, who attempted to reinstall scholarship within the Arboretum landscape during his tenure. Now, after some seventy-five years since the Bussey Institution closed, research and its requisite facilities are unified with the Arboretum's living collections. As a result, visiting scholars from all over the world can work in state-of-the-art laboratories just footsteps from the living collections. Shortly after I began my work at the Arboretum, about a dozen projects occurred in the collections each year. Currently, some seventy-five to one hundred projects use the living collections, landscapes, and environments annually.\nThe Fourth Fifty Years\nThe year 2072 seems so far off. I doubt that I'll be above ground, or if I am, how well I will be able to peruse the collections as they celebrate their bicentennial. Still, if I am around at that time, just a few years shy of my own centennial year, I would like to see the trees from the 1977 expedition to Korea and Japan spreading their branches among the overstory of the collection. I would like to see plants from the Campaign for the Living Collections: some of those trees have recently been released from the nursery and are already taller than me. I'm confident that many will have become standouts the masterpieces of a new generation and subjects of research that we would find impossible to imagine in 2022.\nAnd of course, I would also make my way over to the giant sequoia that overlooks Bussey Brook, checking in to see how it had fared. No doubt, it will have weathered droughts and blizzards, perhaps even a lightning strike due to its ever-increasing height. But I like to imagine it will still be standing, a silent sentry watching over Harvard's tree museum."},{"has_event_date":0,"type":"arnoldia","title":"Saving the World's Threatened Trees","article_sequence":8,"start_page":34,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25754","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170a76b.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Alvarez-Clare, Silvia; Shaw, Kristy; Pocock, Sarah","article_content":"Hiking through the hot, dry canyons at the base of the Sierra La Laguna peaks in Baja California Sur, Mexico, it is impossible to miss the beautiful arroyo oaks (Quercus brandegeei). The trees border the banks of the seasonal streams (or arroyos) like kneeling giants washing their limbs in the refreshing water. What is less obvious is that these represent a relict species that can only be found here, along the riparian zones of the Sierra La Laguna Biosphere Reserve, a biodiversity hotspot with high levels of endemism and great beauty. Each November, the tree canopies fill with elongated acorns that cause a lively commotion as birds, beetles, and rodents frantically eat the fruit on the trees and underneath. Ranchers value the trees too, frequently building corrals under their merciful shade and collecting acorns to feed livestock. However, populations of the arroyo oak are declining. There is no evident seedling regeneration, and the remaining trees are all more than one hundred years old. Until recently, the cause for decline was mostly unknown.\nAcross the globe from the Sierra La Laguna, Mount Mulanje known as the 'island in the sky' rises from the plains of southeastern Malawi with such sheer contrast that it creates its own climate and flora. Best known and most impressive of the forest trees is the cedar that takes its name from these mountains. The Mulanje cedar (Widdringtonia whytei) is highly valued for its durable and fragrant timber, but due to overexploitation and illegal logging, the cedar has reached the point of near extinction. A similar fate is faced by a rare magnolia (Magnolia grandis) found only in the forested limestone mountains of southern China and northern Vietnam. With its large, leathery leaves growing to over a foot in length, this magnolia coexists in tiny forest fragments with other critically endangered species, including the strikingly unique Tonkin snub-nosed monkey. Recruitment of new seedlings is impaired by local agricultural practices in which farmers clear vegetation before planting cardamom and repeatedly weed out the magnolia to maintain their crop. Fewer than three hundred adult trees remain in small, isolated populations.\nThe loss of trees is a global problem. Evidence of declining populations, illegal logging, lack of regeneration, and new pests and diseases has been looming over our heads for decades. Until last fall, however, the complete picture of the status of the planet's tree diversity was unknown. The State of the World's Trees, published in September 2021, shares the results of the Global Tree Assessment the first conservation audit of most of the world's nearly sixty thousand species. The results show that 30 percent of all tree species more than 17,500 species are threatened with extinction. That's more than double the total combined number of globally threatened mammals, birds, reptiles, and amphibians.\nThe Global Tree Assessment also reveals that at least 142 tree species are recorded as extinct. Losing even a single species can have severe consequences for an ecosystem. As primary producers at the base of the food chain, plants, including trees, are the building blocks of ecosystems essential to all life on this planet. Myriad species of plants, animals, and fungi are intrinsically linked to trees, often interacting within complex and fascinating relationships that both parties depend on for survival. In addition, individual tree species play numerous economic, ecological, and cultural roles. We depend on trees in our everyday lives they provide us with food, timber, and medicine. According to the assessment, at least one in five tree species has a recorded human use, and many have a variety of different uses. While the challenges and scale of the problem in maintaining tree species diversity are significant, we can do something about it.\nA Global Campaign\nThe State of the World's Trees is a sobering reminder that trees need our help. The Global Trees Campaign is coled by Botanic Gardens Conservation International (BGCI) and Fauna & Flora International (FFI). Through this effort, researchers, conservationists, and on-the-ground partners have been working together since 1999 to reduce threats and secure or recover target populations of threatened tree species through in situ action. Since its establishment, the campaign has worked to conserve over four hundred threatened tree species in more than fifty countries, and the team has trained more than ten thousand people in tree conservation skills.\nBotanic gardens and arboreta have been vital partners in this effort. Since 2017, for example, The Morton Arboretum, near Chicago, has led a Global Trees Campaign project that aims to safeguard the arroyo oak (Quercus brandegeei) of Baja California Sur. Researchers collected genetic, phenological, and ecological data on this endangered species to explore the causes of decline and identify conservation and management actions needed to save it from extinction. The team established fenced exclosures to quantify the effect of grazing and trampling by free-roaming livestock on seedling survival and growth. They found that cattle and goats eat the seedlings while pigs eat the acorns a combination that prevents any natural regeneration from occurring. To combat these threats, Mexican scientists, land managers, ranchers, and international experts are working together to implement a management plan for this species. Among their actions, the team has conducted plantings within fenced areas to boost population recovery; they have encouraged ranchers to adopt oak seedlings and plant them within their fenced gardens; and they have worked with land managers to establish larger grazing-free zones within the reserve.\nAs illustrated by the work safeguarding the arroyo oak, effective conservation should be informed by accurate baseline information, including a thorough understanding of the species biology, specific threats, and potential actions to mitigate and reverse the decline. Scientific research is one of the cornerstones of the Global Trees Campaign. Once the baseline information is gathered, tree conservationists must develop a plan to improve the success of the interventions. The planning can prioritize individual species, like the arroyo oak or the Mulanje cedar, or larger groups of tree species present in the same area or experiencing similar threats.\nIn Kenya, for instance, Global Trees Campaign partners collaborated with the Kenya Forest Service and the Conservation Planning Specialist Group (part of the International Union for Conservation of Nature) to organize a series of online workshops focused on planning conservation action for Kenya's threatened trees. The workshops brought together key stakeholders to evaluate the results of an analysis for Kenya's more than 140 threatened tree species. This effort helped prioritize sites for conservation by grouping threatened species that are likely to benefit from the same conservation activities. During these workshops, the participants developed a joint vision statement and goals, and they identified actions at national and regional levels. The Global Trees Campaign plans to continue using this larger-scale approach in the future, maximizing efforts and often achieving more cost-effective results than approaches focused on individual species.\nComprehensive Information\nBefore the State of the World's Trees was published, comprehensive information was lacking on which tree species are threatened with extinction and where conservation efforts should be directed. Some assessments were available on the International Union for Conservation of Nature's Red List of Threatened Species and national Red List publications. Still, the information was not easily accessible, and the scale of the problem was unknown. To produce a global overview of the conservation status of trees, the Global Trees Assessment team collated existing assessments, and each species was assigned one of six risk categories: extinct, threatened, possibly threatened, not threatened, data deficient, and not evaluated. Although this effort alone was an enormous task that took more than five years and five hundred contributors, it also revealed the information gaps regarding many tree species. In the report, well over seven thousand species were classified as data deficient, meaning there wasn't enough information for an assessment. Moreover, assessments for many little-known tree species are often based on historic herbarium records that may misrepresent recent changes in land use or loss of populations. Further survey work is therefore required.\nThe information from the Global Tree Assessment can be accessed online via a public web platform, the GlobalTree Portal. The portal highlights the scale of the problem and provides information on the numbers of species found in at least one protected area (as well as species not represented in any protected areas). The portal also shows which species are present in, or absent from, ex situ collections, such as botanical gardens and seed banks. According to the GlobalTree Portal, approximately 56 percent of threatened tree species occur in at least one protected area, and 21 percent are maintained in botanic gardens or seed banks. Another online tool, Conservation Tracker, provides real-time information on who is taking conservation action for which species. These tools will be updated regularly, helping to guide ongoing conservation efforts. The idea is that on-the-ground efforts, such as Global Tree Campaign projects, will use this information and contribute new data as they evolve, creating an information feedback loop that will result in effective conservation actions.\nTargeted Action\nAccording to the Global Tree Assessment report, agriculture and logging are the leading threats to trees globally. When managed effectively, protected areas can provide vital protection against this kind of habitat loss, but in some cases, ecological constraints and threats within protected areas can still prevent or limit regeneration. For instance, even though the arroyo oak occurs within the Sierra La Laguna Biosphere Reserve, natural regeneration has been impossible due to grazing. Tree conservationists must therefore identify and remove barriers to natural regeneration, although additional interventions may be necessary for many species, such as those with extremely small populations. In such cases, planting can be an essential strategy to increase population numbers or reintroduce a species.\nIn the case of Magnolia grandis, with a global population totaling fewer than three hundred adult trees, targeted action was needed to ensure the future of the species. Since 2013, as part of the Global Trees Campaign, FFI has developed an outreach program with local cardamom growers at Tung Vai Watershed Protection Area in Vietnam. These efforts are paying off, with local cardamom farmers now willingly maintaining M. grandis seedlings, indicating a shift in attitudes and behavior towards this species. Over the same period, regular community monitoring and patrolling to protect trees from logging was introduced, resulting in no felling or damage to M. grandis individuals at Tung Vai since 2017. In addition, local communities have adopted fuel-efficient stoves, reducing pressure for firewood. Given the low number of individual trees in the original populations, tree conservationists are conducting booster plantings using nursery-grown seedlings. Natural regeneration of M. grandis is now occurring in other areas of the forest where previously there was none, indicating that recovery work over the last eight years has been successful.\nFor the Mulanje cedar (Widdringtonia whytei) from Malawi, illegal logging was so intense that it removed the natural seed source from the mountain, and increased man-made fires impeded recruitment of remaining seedlings and young trees. As part of a campaign project led by Mulanje Mountain Conservation Trust, the Forestry Research Institute of Malawi, and BGCI, staff set up eight community nurseries around Mount Mulanje with more than eighty community members who had been taught to propagate the Mulanje cedar. Over four hundred thousand seedlings were purchased from community nurseries and planted by local people, providing employment opportunities and vital income. Restoration experts from the Ecological Restoration Alliance of Botanic Gardens are also helping to improve planting practices so that more trees survive and grow better. An extensive network of firebreaks is maintained on the mountain to protect planted seedlings.\nFurthermore, international trade of the Mulanje cedar was restricted when the Convention on International Trade in Endangered Species of Wild Fauna and Flora (a multinational agreement often known as CITES) included the cedar on its list of species that are potentially threatened with extinction. Alternative sustainable uses of cedar are being investigated that could provide additional benefits to local people. Essential oils can be produced from the tree's wood and leaves, and researchers have investigated the components of this oil to identify commercial uses, like soaps. Communities around Mount Mulanje have planted Mulanje cedar hedges from which essential oil can be extracted, and distillation equipment and training are currently being provided. This effort offers local communities alternative incomes from the Mulanje cedar that don't damage Mount Mulanje or its plant resources. The conservation team also planted ex situ trial plots and woodlots elsewhere in Malawi. These actions aim to ensure the planted trees on the mountain remain safe for the long term.\nWhatever approach is taken to reduce threats, improve natural regeneration, or restore populations of the tree species, the full engagement and participation of local stakeholders is key to the success of all tree conservation initiatives. This ensures that the approach is appropriate to the local context, has local ownership and support, and is more likely to achieve a lasting impact.\nThreatened Trees in Restoration\nTrees capture carbon from the atmosphere a fact that has drawn increasing interest given that runaway levels of carbon dioxide are a significant driver of climate change. And trees are also essential components of many habitat-restoration projects. As a result, governments and organizations around the world are investing in large-scale tree planting. These tree-planting pledges and restoration projects provide an opportunity to deliver on conservation goals by incorporating threatened species into the planting plan. However, this opportunity is often missed; many tree-planting projects focus only on exotic species or, even in the case of restoration plantings, only a small number of native species.\nAt Jardim Bot\u00e2nico Ararib\u00e1, in the State of S\u00e3o Paulo, Brazil, a team has been working on a forest restoration project since 1987, intending to restore not only specific plant species but also the entire ecosystem. The efforts at the garden are an exemplar of how threatened species can be incorporated into a successful restoration program. The garden is situated on one of the few remaining fragments of Atlantic Forest. Despite the status of the Atlantic Forest as an important biodiversity hotspot, this forest type is recognized as one of the most degraded ecosystems on the planet. So far, the garden staff has restored about fifty acres (two-thirds of the site). Due to this restoration, headwaters that supply water to Amparo, the closest city, have reappeared. The restored forest protects the riverbanks, preventing silt build-up and protecting the river water.\nThe restoration plantings at Jardim Bot\u00e2nico Ararib\u00e1 feature threatened species, including the endangered brazilwood (Paubrasilia echinata) and another critically endangered species in the legume family, Chloroleucon tortum. The plants for the restoration are grown in partnership with a commercial nursery that also supplies these native tree seedlings to customers for planting in their local area. As a result, the species are becoming part of the local supply chain of native tree species in S\u00e3o Paulo.\nScaling Up Conservation Action\nWith such a vast number of trees at risk of extinction worldwide, a significant scaling up of conservation action is urgently needed. To increase effectiveness and avoid duplication of effort, tree conservationists should mobilize at national levels. It's also crucial to coordinate efforts around specific taxonomic groups, especially genera or families with a high number of threatened species. Species within the same taxonomic group share many characteristics, and they may be subject to the same or similar threats. Therefore, related species are likely to benefit from the same conservation actions.\nBGCI and the botanic garden community have established groups known as Global Conservation Consortia, which are developing comprehensive conservation strategies for highly threatened taxonomic groups identified by the Global Trees Assessment. The consortia aim to coordinate in situ and ex situ conservation efforts and disseminate species recovery knowledge. For example, the Global Conservation Consortium for Oak, led by The Morton Arboretum, mobilizes experts and local partners to conserve oaks, a culturally and economically important taxonomic group that cannot be protected in seed banks. As part of these efforts, the team has organized educational webinars, provided training on seed collection and species propagation, and coordinated regional meetings and workshops focused on filling knowledge gaps for species of conservation concern. To date, Global Conservation Consortia have been developed for six tree groups: oaks (Quercus), magnolias (Magnolia), rhododendrons (Rhododendron), maples (Acer), southern beeches (Nothofagus), and the dipterocarp family (Dipterocarpaceae). These groups include more than eight hundred threatened species, and the model is now also being applied to highly threatened non-tree groups.\nNational coordination of tree conservation efforts is also a valuable approach, as the collaborations in Kenya have demonstrated. The GlobalTree Portal allows tree conservationists to identify countries with high numbers of threatened tree species, especially those with high numbers of threatened endemics. These countries must be priorities for coordinated conservation. Indonesia, for instance, has almost seven hundred threatened tree species, with ongoing habitat- and species-level threats providing little chance for their recovery without dedicated conservation action. While many large-scale conservation programs are dedicated to the country's flagship animals (such as elephants, orangutans, and tigers) or to large areas of high-carbon forest, few initiatives are specifically designed around the conservation needs of individual threatened tree species in situ.\nThrough the Global Trees Campaign, FFI has successfully engaged the Indonesian government in threatened tree conservation. As a first step, FFI established the Indonesian Forum for Threatened Trees, a group of more than seventy members from at least thirty different institutions. The forum convinced the Ministry of Environment and Forestry to consider adding twelve threatened tree species to their list of priority species. So far, one of these trees, a critically endangered dipterocarp known as Vatica javanica ssp. javanica has become legally designated as a National Protected Species. In 2019, the Forum for Threatened Trees and the Indonesian Institute of Sciences published a ten-year national conservation strategy for the twelve priority species. At the same time, FFI also seeks to build capacity for organizations working on threatened trees and inspire new action for priority species.\nMobilizing a Global Community\nIn contrast to the numerous well-known flagship animal species, threatened trees have gained little attention from governments, funders, conservation organizations, the corporate sector, and the public. With 30 percent of tree species shown to be at risk of extinction, this needs to change. Tree conservation requires a concerted response from the global community, with all different regions and sectors engaging and taking action. Botanic gardens and arboreta are in a strategic position at the intersection of research, outreach, and conservation and can play a critical role in safeguarding the world's tree species. The urgency of the situation, however, requires an 'all hands on deck' approach.\nPolicymakers at all levels (global, national, and local) need to incorporate and prioritize threatened trees within legislative frameworks. Intergovernmental and international organizations need to promote and share data from the Global Tree Assessment with their networks and integrate threatened tree conservation into their programs. The corporate sector has an expanded role to play, particularly companies engaged in timber, agriculture, and extractive industries. Land managers, including governments, are key actors in securing critical habitat. Members of the conservation organizations need to prioritize threatened trees within their programs, supporting action on the ground and generating a higher profile for this issue. The tree-planting and habitat-restoration sector have an unrivaled opportunity to integrate threatened trees within their work, contributing significantly to saving species while meeting their other goals. There is a role, too, for the research community. Researchers are necessary for filling information gaps on threatened species and demonstrating the role of tree species diversity in ecosystem resilience. Moreover, there is a need for committed individuals global citizens who advocate on behalf of threatened trees. Now is the time to act."},{"has_event_date":0,"type":"arnoldia","title":"The Making of Arboretum Wespelaar","article_sequence":9,"start_page":44,"end_page":53,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25767","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d14ea36a.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"De Spoelberch, Philippe","article_content":"Whether an arboretum has ten trees or thousands, many of the same management concepts hold true. Yet new collectors often progress through trial and error, as though no one else had gone through the same process. I began raising trees from seed in my garden in the late 1960s. As with many mad collectors (no matter what is being collected), I started the whole thing without much forethought it just began one day. But I kept going and expanded the collection into neighboring woods and meadows. In 2003, I established Arboretum Wespelaar as an independent institution in a small village north of Brussels, Belgium.\nMy family had operated the Artois brewery for generations, so I was fortunate to have the means and the space to begin such a collection. (Artois is now part of Anheuser-Bush InBev, and we are today just long-term family shareholders.) I also had the opportunity of starting early, having good advice from my father, who loved trees, and I was curious and determined to know more. I remember my father kidding me because I did not immediately see the difference between a young beech (Fagus) and hornbeam (Carpinus) or, worse, a spruce (Picea) and Douglas fir (Pseudotsuga). I would not get caught again.\nI returned to Belgium in 1969, after getting a graduate degree in business administration from Columbia University, with 150 seedlings in a big bag. In those days, you could carry about anything on a plane. Most of the seedlings had germinated in a wooden Borden milk box on the terrace of my apartment in New York. I had collected others during a trip to California just before my return. Fifty years later, the redwood (Sequoia sempervirens) from that trip are the tallest and girthiest trees on the estate and arboretum.\nWhile working as a young brewery salesman in my late twenties, I visited dozens of gardens and arboreta around the world. I started buying plants at local nurseries and then European specialist nurseries. The collection spread from the garden around my house (twenty-five acres) into what was to become the arboretum (eventually fifty acres). For the first twenty-five years, I had the help of a single gardener. Now, five-full time gardeners manage the arboretum and the nearby garden at Herkenrode. Over the years, we learned by doing.\nPreparing the Ground\nWhen we began, two kinds of areas were used: meadows and woodlands. Both needed some kind of intervention. I learned this at my expense. Our first foray into the woodlands was done without concern for honey fungus (Armillaria mellea), which causes root rot. As trees were cut to open vistas, we left stumps, and the fungus soon got the best of many choice plants. We did not make this mistake when extending the collections into the old oak wood now in the arboretum: All shrubs and undergrowth were removed with a rotary cutter and uprooted. All deadwood was removed. We did not have additional honey fungus problems, but this exercise did little for the soil structure. It took years before moles arrived, finally suggesting improvements in soil structure and aeration (performed by millions of worms).\nThe old meadows required a different approach. Cattle had trampled and compacted the soils. As a result, it was necessary to plow these areas before planting. In one case, we even allowed a local farmer to grow corn for two seasons. Without soil preparation, the plants sulk, never sending roots beyond the planting hole and eventually drowning there, at least in a flat part of the world like Flanders. After plowing, we created mounds and planted the trees upon them, allowing the water to drain. Initially, it looked as if I was trying to create a minigolf course, but visitors were kind enough to say that the whole thing was not too ridiculous. By now, the result is spectacular. You can easily see that the trees planted on mounds are at an advantage, and the movement in the terrain provides some visual appeal.\nSourcing the Plants\nI have long enjoyed plant propagation. Like many kids, I was fascinated with seeing seeds burst into growth. I was even scolded in school for growing wheat in the inkpot of my desk. The arboretum and the nearby gardens currently contain almost eleven thousand living accessions of woody plants. Of these, 50 percent were raised by us from seed, cuttings, and collected seedlings. Many originated from expeditions to the wild. My first trip was to Nepal in 1975, and successive annual trips (often with the International Dendrology Society) have targeted every possible temperate locale, from California to Hokkaido.\nWhen seeds arrive throughout the autumn and winter, we place them straight into the refrigerator. A numbered label is added to the individual bag and accompanies the seed through subsequent steps. The label is essential. (It is embarrassing to admit that you do not remember the origin of a beautiful plant.) The seed lots accumulate until March, when they are sown in pots. Of course, many seeds could be sown outside when they arrive (the cold, moist winter conditions are generally suitable for this), but mice will always find them and have a feast. Ungerminated seed pots should be allowed to go through another winter, because belated surprises can always be expected. Seedlings are repotted when big enough to withstand the shock (two or four true leaves above the cotyledon) but basically when we have the time. Seedlings can stay crowded in a pot for many months.\nAs a precaution, always split a collection of rare seeds into several lots and treat each set differently. Some twenty years ago, I received a hundred seeds of a recently discovered species of magnolia (Magnolia decidua, then known as Manglietia decidua) from China. I kept fifty seeds and distributed the others in equal sets to five good propagators and magnolia enthusiasts. One morning, I had a look at my tray and realized that a fungus had killed all fifty seedlings. I was hoping that my five colleagues would have succeeded. One had died; one did not remember receiving the seeds. Of the others, Tom Hudson (of Tregrehan Garden in Cornwall) and Dick Figlar (of the Magnolia Society International) had managed to grow the seedlings and are responsible for all specimens of this species in cultivation, including the one at Arboretum Wespelaar.\nCuttings are collected between the end of May until mid-August. Every time we purchase a plant, we immediately take cuttings, given that cuttings from young plants often root more easily. For example, I took cuttings on a young Magnolia 'Elizabeth' three years in a row; out of five cuttings taken each time on the first, second, and fourth year, we succeeded at propagating five, two, and then none. The winter months are hard for the cuttings; even perfectly rooted cuttings will decay under the attack of fungi. Healthy white roots go brown, and the base of the unhardened cuttings does too; the cutting dries up. We have not been very good at keeping our cuttings growing, but these losses can be a relief. We still end up with too many plants: some five hundred cuttings and seedlings every year, which will have to be looked after for another three to ten years.\nSmall seedlings can be collected along roads and edges of woodlands. These will travel well if kept in relatively dry moss, packed in plastic bags or plastic water bottles. (Obviously, you must be respectful of rules and legal restrictions.) We also purchase plants, mainly in pots. The smaller, the better. I have had much disappointment with large plants. Small plants are, of course, cheaper and can be grown to a good size in one of our nurseries until ready for final planting in the arboretum.\nPlanting the Landscape\nWe have used three temporary nurseries around the garden and arboretum. Good woodland soil and shade from large trees provide the ideal growing conditions for our small plants and seedlings. It is ideal to observe your plants until they have suffered a bad winter. It gives you the time to decide where to plant them. They will transplant with a good lump of soil (unlike the miserable peat ball with circling roots that you find at the average garden center). We have seldom failed in transplanting a young tree or shrub raised in these woodland nurseries. On the other hand, we have lost many plants in the first few years in these sites. But better there than in the grounds after an expensive effort at planting!\nWe rarely place a plant directly into its final location. Most spend as long as five years in the nurseries. Few people like the idea; it seems like double work. But I consider not taking this intermediate step to be a grave mistake. Many recently acquired plants will die, and given this reality, I like them to die in the nursery. I have often thought it would have been much better to collect art of any kind and, like a dendrologist, throw two-thirds of the collection away and enjoy the remaining successes. At least, works of art generally gain value over time, whereas aging trees become an expensive problem.\nWhen it comes time for siting the plants, we use a homegrown method involving playing cards. I do not know who came up with the idea, but we have used it for fifty years. We staple two sets of plasticized playing cards (reds and blues) onto plants in the nursery. The identity of the plant and its card is written up on a special form. A corresponding set of playing cards is placed on 104 bamboo sticks, which are reused for several years. We then take a walk through the grounds, staking locations for each of the plants. We aim to get rid of all the bamboo stakes while trying to remain intelligent and effective and still get home in time for dinner. It takes us, in general, up to five hours to place two sets of cards. Of course, we could write the plant's name on the stick, but it is much easier to spot the cards from a distance.\nWe often situate the plants in taxonomic groupings. So, when we're placing the bamboo stakes, we first attempt to place a viburnum, for instance, within the viburnum section. If there is no space left, we find room elsewhere. Obviously, you must know what condition the plant enjoys, how big it will become, and so forth. One becomes better at this with time, but the proper planting distance is always a terrible illusion. Someone once pointed out that when there was a gap between two trees and you add a young tree between them, you end up with two additional gaps. I must admit that I have found myself planting two new trees in such spaces. Discipline is essential.\nLarge trees should be planted at least fifty feet apart, yet we have many at half that distance. We will remove one of them in due course. Trees should not be planted near the edge of a woodland, or they will grow slanted. Likewise, groups of three an arrangement beloved by landscape architects should be avoided as none of the three will end up as a balanced specimen. (This is not a problem for shrubs and small trees.) These conservative approaches will make your arboretum look rather dull for many years, so you have to suffer the irony of friends and guests. Most do not understand what is going on. I like to think that I do not need to see my trees in old age; I know what they will look like. Other plant collectors are more impatient.\nCataloguing and Labeling\nWhen beginning a catalogue for a plant collection, it is a good idea to think carefully over what software to use and then leverage its capabilities to the greatest extent. These days, you may want to consider using relational database software, but a single spreadsheet can be equally effective. Take some time to sit down and think over the structure. Some curators will suggest that at least twenty fields are necessary, but I recommend a minimum of six: accession number, name, landscape location, source, date planted, and condition. Most people will also want a field for any supplementary information. The printed catalogue at Arboretum Wespelaar presently uses nine fields, out of some twenty in our Access database.\nAccession numbers are a difficult concept for beginning dendrologists. I do not know why. An accession number is no more than a simple and unique sequential number given to each plant that comes into the collection. You can give the same number to several plants provided they are from the same source, same age, and in the same location. But otherwise, give them unique sequential numbers, or you will soon regret it. Further, there is no reason to include the date within the accession number.\nOf course, everyone will want to know the age of a plant. Most curators will include the date of planting on the label, which is a good idea. But we made the mistake of including the date as part of the accession number: Our first plant bears accession number 66001, which means it was the first accession in 1966. This system was useful until the year 1999. With the millennium, we got in trouble, as the first plant of the new millennium was 00001. And it shows up first in any numbered list. We had to add two digits for the sequential listing.\nWhen it comes to the name, it is best to refer to a single accepted list, thereby avoiding spelling errors and nomenclatural issues. The Royal Horticultural Society's Plant Finder is probably the only document to be sufficiently comprehensive and regularly updated. Synonyms and taxonomic changes of names are clearly indicated in annual updates. It even includes cultivated varieties. Still, if you specialize in a certain taxon (like magnolias), you may want to use a recent monograph on that group.\nOnce this record-keeping is complete, then comes labeling the curator's nightmare. I have always had an average memory and have not relied on it to know anything. This is probably why I have been so determined to make sure that our plants are properly labeled. Our labels include the name and accession number and are made on a thick ribbon of white PET plastic cut to length and engraved with an automatic engraving machine (a Gravograph). Labels are inexpensive: we estimate that it cost us one euro to make a label with a reasonably long name.\nLabeling problems, however, are never far away. I learned plants while visiting arboreta and botanical gardens all over the world. As I explored these collections on my own, I would go to a plant, take a picture, and then search for a name. I would be exasperated if I did not find a label and sometimes astonished at the number of wrongly labeled plants I encountered. Even so, at Arboretum Wespelaar, one of our members on a study day was surprised to find a label stating Abies rufinerve on a new maple accession. (Abies, of course, is the genus of fir trees the tag should have read Acer rufinerve.) So problems occur even in the best houses.\nChange in the Collection\nArboretum Wespelaar, like any plant collection, is in a constant state of evolution. Not only do plants grow and die, but interests and goals shift as well, changing the landscape over time. Although I fell in love with conifers initially (my first plant was a white fir, Abies concolor, accessioned in 1966), I soon switched to deciduous trees, particularly maples. Around 1969, I went to a local nursery that had a seemingly good catalogue and proudly ordered one of each maple on their list. I soon found that my collection of some twenty maples was far from what the world had to offer. Would I have given up if I had realized that there were more than 120 species, along with hundreds of hybrids and cultivars? My subsequent loves were rhododendrons, magnolias, and stewartias, as proven by the number of those plants in the collection. Today, the team at the arboretum aims to acquire all of the main species in all important genera and in particular plants from wild origin.\nIt is clear that gardens, if well-curated, can contribute to the maintenance of biodiversity. The Franklin tree (Franklinia alatamaha) is a memorable example: although it went extinct in the wild in the early 1800s, the species survived in Bartram's Garden in Philadelphia. While conservation is an additional objective of Arboretum Wespelaar, our primary purpose is to ensure that people can study and learn to love plants. We have no shop, no cafeteria, and nothing for children. Dogs and joggers are not welcome. The result is that our visitors actually look at labels and take notes. I have always intended that the garden and then the arboretum should be open to the public, recognizing that I have benefited from the generosity of botanists, plant collectors, and gardeners who have opened their collections to me. In turn, it's my pleasure to welcome others and inspire them to see and know plants.\nOnce a year, usually in November, we have a difficult day when we deaccession trees, removing them from the collection. This year, we will likely deaccession around fifty plants. These are painful choices but very necessary. We have planted too much with the knowledge that we would have failures and that others wouldn't last. I am adamant that as many as possible of our trees should have lower branches on half of the crown. In due course, aesthetic considerations will always rule above other imperatives. Within a changing collection, it is always nice to have too many good things."},{"has_event_date":0,"type":"arnoldia","title":"A Time for Trees, A Time for Arboreta","article_sequence":10,"start_page":54,"end_page":57,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25764","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170856b.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Donnelly, Gerard","article_content":"Planting and watching trees grow takes time. A long time. The lifespan of a tree may be the equivalent of multiple human generations. This is the good and patient work of arboreta, which requires considerable time horizons to achieve many of their purposes. At The Morton Arboretum, in Lisle, Illinois, we call this 'tree time.' The time required to establish, test, and evaluate tree collections and develop beautiful, planted landscapes that inspire people's interest and appreciation is such that only long-term, multigenerational organizations like arboreta can undertake them.\nGiven these timescales, I like to say that it is good to be old if you are an arboretum. This year, The Morton Arboretum is celebrating its centennial year, having been established by Joy Morton in 1922. Morton had been encouraged and advised by Charles Sprague Sargent of the then-fifty-year-old Arnold Arboretum of Harvard University. The Arnold Arboretum is celebrating its sesquicentennial year, founded in 1872, the same year Morton's father, J. Sterling Morton, established Arbor Day in Nebraska. The Arbor Day Foundation, created fifty years ago, in 1972, upon the Morton-family legacy of planting trees, is advocating for tree planting on the occasion of its anniversary with a theme of 'A Time for Trees.'\nThe time for trees has arrived. There has never been a time when recognition of the value of trees and tree planting was greater than it is today. Trees are being planted globally at scale to sequester carbon and cool the planet. There is widespread appreciation for the ecosystem services that trees provide in urban areas by filtering air pollution, cooling hot cities, and mitigating stormwater pulses. Numerous scientific studies show how trees contribute to human health and well-being.\nYet time has not been good for trees over the past 50, 100, and 150 years. Burgeoning human activities have drastically reduced the size and health of the world's forests as well as the diversity of trees and myriad other organisms that depend on them. In addition, climate change is already impacting trees through altered weather patterns, violent storms and floods, drought, and ravaging forest fires. Trees long-lived, stationary organisms are highly susceptible to climate change because growing conditions are changing at rates that can stress and exceed tolerances and adaptability within their lifetime.\nHow serious is the threat? The recent State of the World's Trees report by Botanic Gardens Conservation International (BGCI) is alarming. Based on the organization's Global Trees Assessment involving contributions from arboreta across the globe, including The Morton Arboretum, the report documented that 30 percent of the 58,497 known tree species in the world are threatened with extinction.\nWith the majority of the world's population now living in cities, urban forests are recognized as key assets to ensure healthful, sustainable, and climate-resilient communities. However, urban centers are challenging settings for trees to grow in and survive, let alone flourish and contribute their full complement of benefits to people, communities, and the environment. Also, trees and their benefits are not equitably distributed across urban landscapes they often reflect the disparities of resources and human demographics.\nClimate change, tree extinction, tree planting, urban forestry, and environmental justice are significant challenges that all arboreta can play a key role in addressing. But the magnitude of these issues requires the power of coordinated collaboration to have a meaningful impact. No single arboretum can do it alone.\nFor this reason and others, ten years ago, The Morton Arboretum established ArbNet as a global network of arboreta. By working together, arboreta can be better equipped to champion the cause of trees. ArbNet has identified more than 2,100 arboreta in 133 countries, all of which have a common purpose of collecting and showcasing the diversity of trees and promoting their planting and conservation.\nArbNet offers an arboretum accreditation program that recognizes standards of professional practice at four different levels of institutional capacity, encouraging the achievement of higher levels of development over time. Lockerly Arboretum in Milledgeville, Georgia, provides a good example. Initially accredited at level two in 2017, Lockerly used ArbNet accreditation standards to set development goals, including the creation of a new horticultural internship program and expanding participation in scientific research. Upon meeting these goals, Lockerly achieved level-three accreditation in 2021. ArbNet helps member institutions exchange information, expertise, and models that others can use or adapt for their purposes.\nClimate change threatens trees as well as the arboreta that maintain living collections of them. Arboreta need to conduct tree performance evaluations and risk assessments to prepare for predicted changes in growing conditions. We also need adaptation strategies that include relocating species, varieties, or specimens to arboreta with more suitable future growing conditions. ArbNet can play a key role in this. Rather than have such exchanges handled variably on a case-by-case basis, an organized system and standardized process are needed to optimize these adaptive plans. The Morton Arboretum envisions a coordinated climate adaptation strategy and program for trees among the arboreta and tree-focused gardens in North America.\nArbNet's interactive network also provides an opportunity to test tree science questions using a 'common garden' approach at arboreta in different growing zones and environmental conditions. One example of this approach is a North Dakota State University study to evaluate adaptive variation among sets of genetically identical poplars (Populus) growing at eighteen arboretum and university sites across the United States (including the Lockerly Arboretum). Researchers are using whole-genome sequencing and climate modeling to predict how plants will respond to different climate conditions in the future and inform management approaches to build climate resiliency.\nTo halt the extinction of threatened tree species, arboreta must commit institutional resources and staff expertise. We must coordinate with one another on targeted tree conservation efforts, including through programs like the Global Conservation Consortia organized by BGCI. A prominent example is the Global Conservation Consortium for Oak led by The Morton Arboretum in collaboration with BGCI and dozens of arboreta and other partners involved in oak conservation. No single arboretum or garden can or should conserve all the world's threatened oak species, so a coordinated, global effort is needed. As part of these efforts, The Morton Arboretum is establishing conservation groves on-site for two threatened species from the southeastern United States: Georgia oak (Quercus georgiana) and maple-leaved oak (Quercus acerifolia). Over tree time in 50, 100, or 150 years curators will use these collections to ensure that the species are safeguarded from extinction risks, and researchers will study what can be done to help them survive in nature.\nTree planting has risen to the forefront as a solution to blunt global climate change, given the ability of trees to sequester carbon from the atmosphere. Yet large-scale tree plantings for carbon sequestration often take the form of low-diversity tree plantations or forestry plantings that do nothing to protect tree biodiversity; they may even diminish it. Arboreta must lend their expertise in tree diversity, planting, and horticulture to improve approaches for carbon-focused tree planting and reforestation efforts. A new global biodiversity standard for large-scale tree plantings being introduced by BGCI will position arboreta and other botanical gardens as key resources to achieve these essential outcomes, ensuring effective carbon capture in addition to not at the expense of biodiversity conservation.\nArboreta also have an important role to play in supporting objectives to plant trees in urban environments to ameliorate heat, filter pollutants, mitigate stormwater flooding, and lower energy costs. Urban forests also add beauty and improve social cohesion, human health, and well-being. Arboreta know how to cultivate trees in designed and managed landscapes, but they must assert their involvement and influence with municipal planners, engineers, and landscape architects to enhance opportunities to develop healthy and sustainable urban tree canopies.\nWhen arboreta partner with community organizations and local government agencies, they can play a meaningful role in addressing the disparities in people's access to the environmental and health-related benefits of a thriving urban forest. Although this issue was not at the forefront of efforts by arboreta or botanical gardens fifty or one hundred years ago, arboreta should now actively seek funding (or commit their own resources) for equity-focused tree plantings that engage residents in participatory planning and provide training for tree planting and care. Arboreta can partner with tree nurseries and growers to provide not only the diversity of suitable trees needed for urban conditions but also at the sizes that can be managed in community and volunteer planting efforts.\nFurthermore, and aligned with the goal to engage and serve a broader spectrum of the public, arboreta must actively foster and support career paths associated with the work of an arboretum to new and different groups of people. Only with a diversified pipeline of tree experts, curators, scientists, horticulturists, conservationists, and educators will arboreta fully serve the public good.\nArboreta, with their beautiful trees and landscapes, attract a substantial public audience and provide immersive experiences and learning moments about the value of trees and nature. These are opportunities to register tree time the time it takes for a tree to reach its full potential over 50, 100, or even 150 years. These long timelines require commitments to tree planting for future generations, sustained efforts to protect them and their growing environment, and actions to address climate change and other tree threats.\nThe grand challenges of our time related to trees require arboreta and tree-focused botanical gardens to collaborate actively. Together, these institutions can achieve more meaningful and successful impacts, engaging their vast collective audience to encourage people to plant and advocate for trees and a more sustainable world. The year 2022 is certainly a time for trees and for arboreta."},{"has_event_date":0,"type":"arnoldia","title":"The Strangeness of Trees","article_sequence":11,"start_page":58,"end_page":59,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25763","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d1708528.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"The general form of a tree trunk, branches, twigs, leaves is so commonplace as to be completely unremarkable. Trees inhabit spaces that most of us experience daily, and, in fact, they often create those spaces. A low, widespread, and rather twisting elm stretches its branches over the patio of a cafe, not far from my apartment. It forms an enchanted ceiling, especially in the spring, when the samaras alight in the branches. Any tree we encounter is likewise defining its space. We move around them, beneath them, and sometimes even upon them. We're so familiar with trees that, for some of us, they feature in our earliest memories. In my case, it was a ten-foot-tall apple tree in a neighbor's backyard. (I filled a bucket with the forbidden fruit and was ordered to return it with an apology.) For Emanuele Coccia, an associate professor at the \u0192cole des Hautes \u0192tudes en Sciences Sociales, it was a trio of Italian umbrella pines viewed from the balcony of his childhood bedroom. He calls them his 'first image of the world.'\nCoccia recounts this memory in Trees, a book designed for an exhibit of the same name at Fondation Cartier, a contemporary art museum in Paris. The large-format book, published in an English translation, is the kind that you might see stacked on a coffee table in a furniture catalogue. It's filled with almost five hundred images, including field sketches, conceptual paintings, and film stills. Often, when parsing meaning from an artistic depiction of a tree, we turn to a standard suite of metaphors. We see the ancient oak, gnarled and twisted, as a symbol of endurance and solidity. We see a small tree growing from broken concrete as a reminder of perseverance. Scholars might examine specific depictions through the lens of post-colonial studies or otherwise. Yet many of the writers and artists who contribute to Trees suggest that, first and foremost, we must acknowledge trees' status as living beings, reconsidering the strangeness of their too-familiar forms.\nStefano Mancuso, the Italian biologist who is a prominent figure in the controversial field of 'plant intelligence,' leads this charge, pointing out the bizarro ingenuity of plant life. 'Like the negative of a photo, what is white in the animal world is black in the plant world,' he writes. 'Organisms that are so different from us that, as far as we are concerned, they may as well be aliens that evolved on a different planet.' Mancuso enumerates many of the differences between the lifestyles of plants and animals, including differences pertaining to movement, of course, and our inverse needs for carbon dioxide and oxygen. He emphasizes one difference as especially noteworthy: the distribution of specialized functions. While almost all animals have organs that cannot be separated from the rest of the body, plants spread these functions across their form in repeating modules. Plants, for instance, respire without organs that resemble lungs. They digest food without anything that resembles a stomach. Given this functional distribution, a Kentucky coffeetree can lose a large branch from a lightning strike (another one of my early childhood memories) yet retain its ability to produce the organic compounds needed to continue living.\nThis phenomenon of distribution, Mancuso suggests, can cause us to discount the liveliness of plants. We recognize that plants are living organisms, yet we see little of ourselves in their structure. Although we know that plants die, many of us aren't exactly sure what it means for them to be alive. Distribution, we come to recognize, is fundamental to the forms featured in Trees.\nAmong the most maximalist works in the book are Luiz Zerbini's large-scale paintings that situate trees within a jumble of urban textures. Zerbini's Mam\u00e3o Manilha shows a potted papaya (Carica papaya) growing alongside several bromeliads. Two papaya leaves sag along its trunk, preparing to join another that has already dropped to the ground. Above them, a bird opens one of several fruits, revealing the orange flesh and black seeds within, and above that, white flowers appear in large, loose clusters. The painting not only captures the modular form of the plant each leaf, each fruit, ultimately destined to be shed it also captures how this disposability becomes central to a web of other biotic interactions. The pot suggests that a human had grown the papaya in anticipation of the fruit, yet, in a war of attractions, a bird won the harvest. A series of leaf scars along the papaya's trunk also reminds us of the seasons of growth and disposal that have led to this moment. The painting is a composite an imagined place yet the plant seems to be a singular individual, forging an existence that is less than glamorous but nonetheless alive.\nThe book also includes works by Indigenous artists from several regions in South America, including the Gran Chaco, the semiarid plain that sprawls between the Paraguay River and the Andes. The works from this region are ink and paper drawings, and almost all capture interactions among trees and other organisms. A fascinating untitled work by Eurides Asque G\u2014mez shows lines of leafcutter ants trailing into their volcanic burrows carrying leaves of algarrobo trees (Prosopis nigra). The ants, in turn, are shown being picked off by partridges. According to the artist, who is quoted in an essay by Ursula and Verena Regehr, the partridges nest in grasses between the algarrobo, knowing that the ants are partial to the young leaves. Meanwhile, an ovenbird has built its nest in one tree, and birds perch on the branches of another. In this way, G\u2014mez showcases not only the modular, throw-away nature of the trees' emerging leaves a solution to being immobile targets for predators but the way that their modular structures become essential to other organisms.\nMoreover, Trees is a testament to the ways these omnipresent forms shape the lives of humans. G\u2014mez and other artists often include people in the web of arboreal interactions depicted in their art. An atmospheric scientist, Abigail Swann, describes how trees influence climate, choreographing weather patterns a fact among many in the book that reminds us that our disregard for the imperiled state of trees may precipitate our own demise. Yet, on a personal level, the artists and essayists are, themselves, residing among trees, sometimes building livelihoods around their forms. The ensemble of individuals includes landscape designers, a mathematician, the film director Agn\u008fs Varda, the American artist Charles Gaines, and many others.\nYet it is the botanist Francis Hall\u00e9 whose lifelong engagement with trees is most clearly documented in the pages. Hall\u00e9 offers forth drawings from field notebooks, prepared in rainforests around the world: Sri Lanka, C\u00f4te d'Ivoire, Peru, and elsewhere. In some sense, these field sketches represent the leaves of Hall\u00e9's career, collected and pressed within the covers of dozens of notebooks that he has labeled by date and location. 'You quickly realize that the shape of a tree, even when young is never random,' Hall\u00e9 says in an interview with Coccia. 'Each species has its own 'architectural model,' that is, a tree's growth and development follow a genetic program.'\nHall\u00e9's drawings endeavor to capture these unique forms. Among his most impressive works is a large drawing on tracing paper titled Forest Profile, which depicts dozens of trees growing in relationship with one another in French Guiana. He provides two views of the forest: from the side (a cross-section that shows the complex layering of tree canopies) and from above (showing the locations of the tree trunks and the spread of their branches). Even in this schematic form, Hall\u00e9 captures each individual's species-specific, non-random shape. His empirical approach seems like it would produce results that are more like traditional scientific illustrations, often beautiful but unsurprising. Yet through his careful attention to detail, and the disambiguation of these overlapping forms, Hall\u00e9 captures what many of the other artists in Trees likewise reveal: the strange reality of the still lives of trees."},{"has_event_date":0,"type":"arnoldia","title":"Gardens for All","article_sequence":12,"start_page":60,"end_page":63,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25765","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170896e.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Mack, MaryLynn","article_content":"As an African American woman who has worked in public gardens for the past eighteen years, I often experienced firsthand the need for greater diversity. The lack of inclusion in the workspace is not an issue exclusive to public gardens, but it should be noted that many public gardens in the United States were founded by white people and many are primarily staffed by white people, despite being located in communities of color. Like many of my friends and colleagues in other industries, I was often asked to be the representative for all people of color when discussing inclusion, diversity, equity, and access (a set of issues often known as IDEA). Being the 'only' in a room was disconcerting, but it also gave me access and opportunity to speak on important matters and empowered me to do my own self-reflection, do my own research, and do my best to connect and engage with as many communities as possible.\nOver my years working in gardens, I found myself having conversations with employees at other botanic gardens and arboreta who also served communities not reflected in their boards, staff, and volunteers. I may have been an 'only' in my workspace, but I was far from alone to bring forward the need for change. The American Public Gardens Association (APGA) also had conversations with its members and took the call to action to begin a more authentic discussion about the bias, barriers, and baggage in our industry. APGA is the leading professional organization for the field of public horticulture. Members include more than ten thousand individuals at over six hundred institutions, in all fifty states, the District of Columbia, Canada, and twenty other countries. The organization's primary goals are to encourage best practices, offer educational opportunities, and advocate for members, so this dialogue was a crucial step toward action for public gardens as a whole.\nIn 2016, a group of eleven truth-seekers scheduled a phone meeting to talk about diversity and inclusion. This group wasn't appointed, but we were individuals who had asked questions or nudged the association to 'do something.' We represented generational, ethnic, gender, racial, and sexual-orientation diversity and worked in gardens throughout the United States. Only a handful held a high-level leadership position in their respective gardens. This inaugural group spent the first twenty minutes dissecting the definition of diversity. Through that process of discovery, we unearthed the varying degrees of knowledge, the chasm of feelings and opinions, and a quick understanding of just how different we all felt on how to move forward. While at times uncomfortable, we realized that within that uncomfortable space, we could reflect on our own bias. Thus began a year-long exploration of reading diversity articles, untangling historical perspectives steeped in garden history, and having informal chats about our own experiences while serving public gardens. The work was difficult and sometimes frustrating, without a guidebook of boxes to check.\nIt is important to note that regardless of where gardens and their staff stand in their work towards inclusion and diversity, everyone must start by addressing what they do not know. Starting with a garden's history, for example, gardens should bring to light what the land was before, and who lived on it and cared for it. One resource that is especially helpful when exploring this issue is a book, edited by Duane Blue Spruce and Tanya Thrasher, titled The Land Has Memory: Indigenous Knowledge, Native Landscapes, and the National Museum of the American Indian. It speaks to the rich history and contribution of indigenous people to the land in the Americas and is a thoughtful representation of how traditional Indigenous ways should be put into practice by cultural institutions.\nIncreasing individual knowledge in these and other areas is crucial. This work helps combat the collective unawareness that exists when members of a group believe that others in their group hold comparably more or less extreme attitudes, beliefs, or behaviors. The term 'unawareness' is not meant to disparage the work currently happening in gardens but is a reminder that the work needs to start with recognizing that the struggles of communities of color are not new. Allies must take advantage of resources that include research reports, academic studies, and courageous conversations that bring to light past disparities.\nAfter a year of self and group discovery, the IDEA committee made plans to involve the membership at the APGA conference held in Hamilton, Ontario. Our inaugural session was scheduled for eight in the morning on Saturday, a tough time slot since it was the last day of a week-long conference and the morning after the traditional evening farewell celebration. We were nervous and truly had no idea how our stories and messages would be received. The agenda was informal: committee members had decided to simply introduce the topic of diversity and share personal experiences. With a mere five minutes before the session started, we had to request more chairs the room was already packed. What happened next showed us that public gardens were ready and eager for change.\nIn that crowded room, we had executive directors of large gardens, first-time attendees, educators, gardeners, and outreach coordinators who worked directly in their local communities. We listened, shared personal vulnerabilities, and publicly accepted a challenge to move forward in the work. Many conversations continued in the corridor after the session. We were all so excited, but we all had the question: What in the world needed to happen next?\nInclusion and diversity work is often slower than people might hope for. It takes time to develop authentic relationships, actively listen, and recognize that every public garden has different obstacles to overcome. It also takes time to build trust. Patrick Lencioni, author of Five Dysfunctions of a Team, writes about this, describing team-building steps that also work when creating a more inclusive environment. The fear of being vulnerable is often a barrier when speaking on matters of race, diversity, and equity, yet showing vulnerability builds trust in conversation and in relationships. Asking questions that allow people of color a safe space to share their experiences of microaggressions, gaslighting, and other forms of bias are first steps toward changes needed in the workspace and the garden.\nA few years ago, I had the opportunity to speak on diversity and inclusion at the Botanic Gardens Conservation International Congress in Warsaw, Poland. During the open time for questions, one attendee expressed his concern when broaching a conversation about race with someone in the workplace. This person was afraid of using the wrong words, saying the wrong thing, or inadvertently offending a colleague. We discussed the need to acknowledge your own bias and privilege, but then I ended by stating that you just need to 'step in it.' Not my most eloquent moment, especially since I was attempting to encourage this person to step bravely toward having the conversation rather than becoming immobilized and missing an opportunity to have an authentic exchange. Yet that became my tagline and the start of many meaningful conversations for the duration of the conference. This work is messy, imperfect, wonderful, and needed.\nMany public garden leaders have embraced this need for diversity and inclusion and entered into the work with vulnerability and passion. Brian Vogt has built a framework that infuses IDEA throughout every aspect of Denver Botanic Garden's operation, where he is the chief executive officer. For over ten years, the garden has devoted themselves to IDEA principles with board and staff committees, as well as extensive relationship development resulting in eighty partner organizations. When describing their approach at the garden, Vogt notes how they 'emphasize the power, not the pain, of IDEA work.' Today, their visitors reflect the diversity of their community as does the board itself, which is now 40 percent non-white. These changes have resulted in programming that lifts up diverse voices, exhibits, and communications.\u00caVogt further emphasizes, 'Don't get distracted authentic diversity and inclusion work makes\u00caeverything\u00cabetter.'\nOther gardens initially take an external approach and achieve sustainable results. Bruce Harkey, the president and chief executive officer of Franklin Park Conservatory and Botanical Gardens in Columbus, Ohio, led an effort to improve the quality of life in the community by creating neighborhood-based outreach and educational programming. One recent example is the conservatory's participation in the HeART of the Protest, where the King Arts Complex produced forty-six days of artistic projects to honor the forty-six years of George Floyd's life. Franklin Park Conservatory hosted performances by dancer Candice Igeleke and musician K. Daniel. These events presented new work that focused on telling the story of Black Americans, from slavery to present day. Franklin Park recently added an internal focus: the board, leadership team, and staff work in unison to honestly assess their diversity, equity, and inclusion status. They then set goals and objectives for measurable improvements.\nThese and countless other examples show that our gardens are embracing change. After APGA's initial group session in Hamilton, members expressed a growing interest to hear and do more when it came to IDEA principles and practices. The following year, in 2018, when the IDEA committee presented at the Southern California conference in a capacity-filled ballroom, it became apparent we were more than ready to make inclusion a collective goal. The next year, in Washington, DC, the entire conference theme was Diversity. This resulted in a week-long conference filled with panel discussions, lectures, and facilitated sessions surrounding topics about diversity, equity, accessibility, and inclusion in our gardens and beyond.\nOne key moment happened during our very first IDEA Caf\u017d, a keynote luncheon where a group of esteemed panelists talked about their own experiences in public gardens. One of the panelists was in a wheelchair and needed to use the elevator to get backstage. With mere moments before the group was to take center stage in front of an audience of hundreds, the hotel manager received a radio call that the elevator was stuck with our panelist inside! The situation was rectified but we decided to use what happened as a teaching moment. This was an example of how accessibility issues are always present and can impact a person's experience in significant ways. These shared experiences and conversations inspired us to build systems and best practices for the APGA sustainability index, gather feedback and success stories from gardens, and provide encouragement for those gardens who are just beginning to address these issues.\nI smile recalling Brian Vogt's charge to 'embrace the work of diversity and inclusion joyfully.' It is good advice. While our work with inclusion will never be done, the past two years have taught us that collective resilience and embracing change will sustain us along this journey. As I think about the diversity of plants in my garden, which experience stress and environmental adversity year after year, I'm amazed by how they somehow adapt and persevere through it all. They are resilient, and so are we."},{"has_event_date":0,"type":"arnoldia","title":"Balling and Burlapping","article_sequence":13,"start_page":64,"end_page":64,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25766","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d14ea326.jpg","volume":79,"issue_number":"1","year":2022,"series":null,"season":null,"authors":"Copeland, Chris","article_content":"Transplanting is a delicate process that ideally occurs during dormancy, at the beginning or end of the growing season. In the Arnold Arboretum's nurseries, we use traditional methods to ball and burlap our field-grown trees. To preserve the proper ratio between roots and shoots, we measure the diameter of the trunk: for every inch, we need a minimum root-ball diameter of fifteen inches. Because we cannot input every factor into an equation, we also exercise judgment, accounting for the tree's height and the anatomy of its root system.\nOnce we have determined the diameter, we sever the roots with a sharp spade and excavate a trench. The root ball should be deep enough to ensure that taproots are retained at least 65 percent the diameter. We shave away excess soil to minimize transport weight. The exposed root ball is wrapped with burlap and secured with sisal, using a drum-lacing pattern. We carefully rock the tree, freeing it from the soil below. At this point, the tree is ready to go."},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25759","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d170b726.jpg","title":"2022-79-1","volume":79,"issue_number":"1","year":2022,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Arnoldia Reimagined","article_sequence":1,"start_page":2,"end_page":5,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25736","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25e856a.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"This issue of Arnoldia is devoted primarily to the world of nineteenthcentury horticulture and botany, the milieu that shaped the Arnold Arboretum upon its founding in 1872. Yet, in some sense, the issue also represents the culmination of a twentieth-century vision for the magazine itself. Next year, as part of the Arnold Arboretum's sesquicentennial celebration, Arnoldia will relaunch with a structure and approach that is dynamic and distinctly modern. The magazine will still appear in print every quarter and serve as a definitive source for novel and interdisciplinary research on trees, shrubs, and landscapes. Yet, an updated format will allow for new points of access\u2014new kinds of content. In the context of modern publishing, the production of a magazine like Gardener's Monthly, which began in Philadelphia in 1859, seems almost inconceivable. Its editor, Thomas Meehan, would have exchanged feedback with authors on handwritten manuscripts. That much can be expected. More miraculous was the printing. The final manuscript would have been typeset by hand, each page composed of thousands of individual lead characters. Once a page was complete, a proofreader would review a test copy, marking errors as an assistant read the original manuscript aloud. According to a detailed account of the process for producing Harper's Magazine, outlined in 1865, the initial proofs were often rife with errors. After all, the compositor prepared everything backward, in the inverse of the printed page. After corrections and additional proofing, the process would continue to the individuals responsible for operating the presses, folding machines, and so on\u2014an elaborate, labor-intensive coordination of both mechanical and human power.1 The Arnold Arboretum's first foray into magazine publishing was a monthly titled Garden and Forest. It debuted in 1888, weeks after Gardener's Monthly ended. Charles Sprague Sargent, the first director of the Arboretum, oversaw the magazine for its ten-year run, but the editorial offices were in New York, a few blocks from the printer: Harpers and Brothers. (Harper's Magazine was produced in the same building.) Arnoldia was born as The Bulletin of Popular Information in 1911, and for the next fifty-nine years, the periodical was typeset by hand, using the same basic method employed for Gardener's Monthly. The final person to perform the tedium of creating Arnoldia word by word, line by line was Howard Allgaier, the printer for the Harvard University Botanical Museum. Allgaier began producing the publication in 1933, at the behest of Oakes Ames, the supervisor of the Arnold Arboretum. Ames, a bibliophile, was known to say that \"a botanist's research should be a jewel worthy of a proper setting.\"2 Ames also widened the purview of the Bulletin. For its first two decades, the periodical had focused almost entirely on plants growing at the Arnold Arboretum, but in 1931, the format shifted to standalone, topical articles. Ames wrote several of these, including one on the botanical drawings of John Singer Sargent. Arnoldia Reimagined Jonathan Damery Facing page: In the early 1930s, when Arnoldia was still known as The Bulletin of Popular Information, an interdisciplinary spirit emerged that continues to inspire the magazine today. Blanche Ames provided its first contemporary illustrations. ARCHIVES OF THE ARNOLD ARBORETUM His wife, Blanche Ames, began supplying botanical artwork of her own. The following year, their son coauthored an article about searching for beach plums (Prunus maritima) from an airplane. Authors would follow their wide-ranging lead. The name of the publication changed to Arnoldia in 1941, but otherwise, the structure and general approach remained the same. In 1970, Arnoldia relaunched under the production of a new printer, the Harvard University Printing Office. At least through the end of the decade, Arnoldia was produced on \"hot type\" machines, which meant that the words were input on a keyboard and cast from lead on the spot.3 This mechanical process had emerged almost a century before, but perhaps owing to the relatively simple one-article format of Arnoldia, it had remained feasible for Allgaier to continue setting the type by hand. The change in printers coincided with a dramatic reimagining of Arnoldia\u2014a project overseen by Stephanne Sutton, who took over the publication upon the retirement of Donald Wyman, the editor for twenty-nine years.4 The 1970 redesign was more than a visual makeover; it also brought new storytelling approaches. The 1960s is often considered an era of innovation in magazine publishing. Large general-interest magazines experienced circulation declines, attributed to the rise of television. (For instance, Life, which once claimed to reach the hands of one in four American adults, ceased publication in 1972.) At the same time, special-interest magazines began to proliferate.5 The redesign of In 1970, Arnoldia was reimagined as a special-interest magazine with multiple features per issue. The current logotype of Arnoldia debuted at the end of 1982. ARCHIVES OF THE ARNOLD ARBORETUM Arnoldia 5 Arnoldia firmly repositioned the magazine within this new publishing context. While Arnoldia had long hosted a diverse mix of subjects, authored mainly by horticultural professionals, it would thereafter contain multiple articles per issue and showcase a glossy image on the cover. Over the next five decades, Arnoldia went through several visual updates. Among those milestones: the current logotype and dimensions debuted in 1982, and the first color photographs appeared on the interior pages in 2001. Behind the scenes, the modes of production changed dramatically, but our graphic designer, Andrew Winther, skillfully maintained the visual continuity. He began working on the magazine in 1986, while in the art department at the Office of the University Publisher. At that point, the office used offset lithography, and the printing plates were created from photographic negatives of the text and images. By the early 1990s, Winther began designing the layouts on a computer, and ultimately, every aspect of prepress production has gone digital as well. Despite these changes, the basic architecture introduced in 1970 has endured, with each issue composed primarily of several long-form features. In 2022, when the redesigned Arnoldia launches, the feature articles that have long defined Arnoldia will remain central to each issue. But in the opening pages, we will provide a new, distinctive space for shorter narratives that capture behind-thescenes experiences of working with plants in the twenty-first century. We're also adding space for letters, to foster a public dialogue with you, our readers. In the back, we're creating a department composed of essays and opinions. We'll also incorporate contemporary artwork throughout the magazine, building on the legacy established by Blanche Ames ninety years ago. With the first issue of Garden and Forest, published on February 29, 1888, Sargent and the other creators described their commitment to sharing \"noteworthy discoveries\" in the realm of science and horticultural practice. They promised that the magazine would \"place scientific information clearly and simply before the public, and make available for the instruction of all persons interested in garden plants the conclusions reached by the most trustworthy investigators.\" Articles would cover landscape gardening, forest conservation, entomology, and more. The authors would deal both in history and news. Here, looking into 2022, we're doubling down on these longstanding commitments. Expect the first issue to arrive in March 2022. Notes 1 Guernsey, A. H. 1865, December. Making the magazine. Harper's New Monthly Magazine, 32(187): 1-31. 2 Allgaier, H. J. 1984. The printing shop. Botanical Museum Leaflets, Harvard University, 30(1): 48-50. 3 Ashton, P. S. 1980. The director's report: The Arnold Arboretum during the fiscal year ended June 30, 1980. Arnoldia, 40(6): 238-293. 4 Howard, R. A. 1970. The director's report: The Arnold Arboretum during the fiscal year ended June 30, 1970. Arnoldia, 30(6), 201-250. 5 Abrahamson, D. and Polsgrove, C. 2009. The right niche: Consumer magazines and advertisers. In D. P. Nord, J. S. Rubin, & M. Schudson (Eds.), A history of the book in America: Volume 5: The enduring book, print culture in postwar America (pp. 107-118). University of North Carolina Press. Jonathan Damery is the editor of Arnoldia."},{"has_event_date":0,"type":"arnoldia","title":"The Trees of the Silent Dell","article_sequence":2,"start_page":6,"end_page":7,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25737","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25e896d.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Barnett, David","article_content":"A cemetery, by nature, is a place where the past is always present. On September 1, 2021, I retired from Mount Auburn Cemetery in Cambridge, Massachusetts, on the twenty-eighth anniversary of the day I started employment there. I had arrived in 1993 as the director of horticulture, having a background in public garden management and degrees in horticulture and ecology. At first, I only noticed the natural landscape and the spectacular collection of trees. Mount Auburn, after all, occupies a unique space in the history of American landscape design: It served as inspiration for other pastoral cemeteries in the mid-nineteenth century and, subsequently, for urban green spaces like Central Park and the Emerald Necklace. I didn't initially focus on the monuments and the other \"cemetery\" aspects of Mount Auburn. About two years after my arrival, I gave a tour of Mount Auburn to Richard Harris, my major professor from graduate school at the University of California, Davis, who had authored a textbook on arboriculture. We stopped in Consecration Dell, a natural amphitheater in the center of the cemetery, where paths on the shaded slopes overlook a small pond. I explained that we had just initiated a project to restore this area to the woodland habitat that existed when the cemetery was founded in 1831. In fact, Mount Auburn's first president, Joseph Story, delivered his consecration address in this very location, noting the importance of natural beauty when mourning loved ones. \"What spot,\" he asked, \"can be more appropriate than this, for such a purpose.\" I described how the restoration would require a phased approach to remove all exotic plants, especially invasive species such as Norway maple (Acer platanoides), and replace them entirely with native species of trees, shrubs, and woodland groundcovers. I felt proud to describe to my mentor how the restoration plan would allow me to put into practice ecological management concepts that I had studied in graduate school. We happened to be standing next to a spectacular Japanese stewartia (Stewartia pseudocamellia) planted in 1939. I noted that we would not remove the stewartia just because it was an introduced species, but that, when the stewartia eventually died, we would replace it with a native. I also pointed out that the stewartia had a memorial plaque on it with the name and birth and death dates of a woman who had recently passed away. As we talked, a woman who had been walking nearby came up to introduce herself. She was the daughter of the woman memorialized on the tree plaque. She told me that the family had chosen to purchase the plaque because Consecration Dell was one of her mother's favorite spots. The woman said she visited frequently to think about her mother and thanked me for making Mount Auburn\u2014and Consecration Dell itself\u2014such a beautiful, uplifting, and inspirational place. From that day forward, my relationship with the landscape changed. Talking to the woman beneath the stewartia, I came to understand the significance of Mount Auburn as a cemetery and the importance of serving our \"clients\" with compassion and sensitivity. The entire staff understands this\u2014it is embedded in our culture. My colleagues have all had interactions with visiting family members similar to the one I experienced that day. These encounters motivate us to continue achieving the high standards of maintenance of the grounds\u2014from the trees and gardens to the monuments and other built structures\u2014in order to ensure that Mount Auburn Cemetery remains the beautiful and inspirational place that Joseph Story and the rest of our founders envisioned in 1831. The successful restoration of the native woodland in Consecration Dell over the twenty-five years since that memorable conversation has been one of the highlights of my career. In place The Trees of the Silent Dell David Barnett Consecration Dell represents a nearly two-hundred-year-old vision for the naturalistic landscape at Mount Auburn Cemetery. PHOTO BY THE AUTHOR; MAP FROM HARVARD MAP COLLECTION, HARVARD UNIVERSITY of the Norway maples and other invasive species that we removed, hundreds of native trees and shrubs and thousands of ferns and woodland groundcovers now provide a valuable habitat for the birds, salamanders, and other wildlife residents of Mount Auburn. And yes, the magnificent stewartia remains as well. I like to think that the landscape looks just like \"the hill and the valley, the still, silent dell, and the deep forest\" that Joseph Story described so long ago. David Barnett was appointed president and CEO of Mount Auburn Cemetery in 2008. He retired from that position in 2021 confident that the course has been charted for a bright and successful future as an active cemetery, a historically significant cultural landscape, and a model of environmental stewardship."},{"has_event_date":0,"type":"arnoldia","title":"The Nauvoo Rose on Temple Square","article_sequence":3,"start_page":8,"end_page":9,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25738","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160a325.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Henrichsen, Esther Truitt","article_content":"\"She brought it from Nauvoo, Illinois, to Salt Lake City in a teapot,\" my boss, Peter Lassig, told me. It was the spring of 1980, and we were standing in a quiet corner of Temple Square, in the heart of Salt Lake City. Before us, a small, unglamorous rose was beginning to produce its small, deep-red flowers. Peter had asked me to transplant it to a historic home garden, two blocks away. The rose was growing within a collection of special plants protected by the warmth and shade of a fifteen-foot wall made of adobe and sandstone that surrounds the square. Peter explained that the rose came from a woman named Elizabeth Hubble. \"She walked the thirteen hundred miles from Nauvoo,\" he said, \"but her rose rode in the wagon and was most likely the only luxury she allowed herself.\" Elizabeth was one of seventy thousand Latter-day Saints who made the trek across the plains along the Mormon Trail from 1847 to 1869 before the railroad connected the West to the rest of the continent. Elizabeth was among those who were expelled from their homes in Nauvoo, a city they had built. She would have had little time to dig the plant from her garden, and she made a real commitment to keep it alive for the rest of her journey. She would have watered it from the Platte River in Nebraska, the Sweetwater River in Wyoming, and Emigration Creek as she traveled down into the Salt Lake Valley. As Peter told me about the storied rose that late spring afternoon, we were standing across from the south door of the Assembly Hall, a beautiful, Victorian Gothic building, completed in 1882, that was about to go through an extensive renovation\u2014the reason it was necessary to move the rose. Temple Square is the most visited site in Utah, which is impressive for a state boasting five national parks. Its ten acres are dominated by the large, domed Tabernacle and the Salt Lake Temple, divided by the Center Mall. With a cathedral of fabulous American and European elms (Ulmus americana and U. laevis) overhead, Temple Square has served as one of the great urban spaces in the United States for well over a hundred years. The perimeter wall was built as fortification when Salt Lake City was still wilderness and now provides a peaceful space amid the noise of growing urbanity. The next morning, I took a shovel and a pot to dig the little Nauvoo rose, becoming one more in a line of gardeners who had cared for the plant and its provenance since Elizabeth's family had given it to Temple Square in the 1880s. Peter had been introduced to the rose in 1953, when he was fifteen, by his boss Irvin Nelson. In turn, Irvin had been charged with caring for it by his predecessor, who had gardened at Temple Square since the late 1800s. This location was the second placement for the rose on Temple Square. I was taking it to its first new home in nearly a hundred years. Towering over the rose were three Japanese tree peonies (Paeonia suffruticosa) that were the most tree-like peonies I have ever seen. They had been a gift in the 1930s from Brown Floral, a family-run nursery that is still part of the horticultural fabric of Salt Lake City. Each plant had at least thirty mauve blooms, and they were dug and moved to the garden south of the Temple. Several other plant treasures in this space would also be transplanted. In the spirit of its century of being a repository of gift plants, this garden between the Assembly Hall and the Temple Square wall was where, six years later, I chose to plant the seven-son flower (Heptacodium miconioides). This plant was sent to subscribers of Arnoldia when the story of this newly introduced species was published in the Fall 1986 issue. That Heptacodium grew into a glorious tree that every few years bloomed at the same moment as the monarch butterfly migration from north to south. You could stroll past the tree and be amazed as hundreds of monarchs were startled into the air. It was cut down a few years ago by a gardener who had no knowledge of its history The Nauvoo Rose on Temple Square Esther Truitt Henrichsen HENRICHSEN, E. T. 2021. THE NAUVOO ROSE ON TEMPLE SQUARE. ARNOLDIA, 78(5-6): 8-9 and was cavalier about not wanting to learn from those who had come before. In the process of digging the rose that morning in May 1980, I was horrified when it split in two. But, this became an opportunity. I carried the little plants across the two blocks to the Beehive House, where I was the summer gardener and weed-puller. I planted them on either side of a path that led to a gate in the cobblestone wall. Brigham Young had built the wall in the 1850s around his two side-by-side homes, the Beehive House and the Lion House. The roses flourished there for two decades, until the cobblestone wall suddenly collapsed, killing one of the pair. The other was moved to another part of the Beehive House garden while the wall was being rebuilt and was never moved back. I was concerned for the future of the Nauvoo rose because it was difficult to find anyone in the next generation who was interested, but I eventually took three cuttings and have grown them in my home garden for the past decade. By the time this map of Salt Lake City was published in 1870, the Nauvoo rose had been growing in the community for two decades. The rose can now be found in the gardens of the historic Beehive House, mapped with a number 5. PHOTO BY THE AUTHOR; MAP FROM LIBRARY OF CONGRESS, GEOGRAPHY AND MAP DIVISION I once keyed out the Nauvoo rose and believe it is a Rosa chinensis 'Minima', a variety (formerly known as Rosa indica minima) introduced into cultivation in the early 1800s. It grows about two feet high and two feet wide, and it blooms from spring to fall. In the intense high-desert sunlight of Utah, it prefers growing in a bit of shade. Compared to other roses, the Nauvoo rose may not seem very glamorous. Elizabeth, however, had the imagination to envision her little plant blooming in her new home in the Great Basin. Her descendants who donated the rose and the line of gardeners who cared for it since have all been connected by the love, care, and determination required to let it grow. Esther Truitt Henrichsen is the garden designer at Thanksgiving Point Institute in Lehi, Utah. Previously, after completing a master's in landscape history, she worked for many years as a landscape designer at Temple Square."},{"has_event_date":0,"type":"arnoldia","title":"Five Generations of Russell's Garden Center","article_sequence":4,"start_page":10,"end_page":12,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25739","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160a728.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Russell-Skehan, Elizabeth","article_content":"I felt the presence of the large video camera and mic over my right shoulder as I opened the photo album of Russell's Garden Center from the 1980s. \"There's the four of us,\" I said with a smile to my husband, Tim, who sat next to me. I was referring to a photograph of us with my mom and dad, wearing our teal Russell's shirts and sitting in front of our new sign on Route 20. The highway connects Wayland with Boston, about sixteen miles to the east. Our daughter Genevieve, the movie director, encouraged me to continue. \"Pretend there's no camera or mic here, and just tell me about the five generations of Russell's.\" I began my story, explaining how the business was established in 1876. \"My great-grandfather Samuel Lewis Russell was a butcher,\" I said, \"and his original store was called Russell's Provisions.\" He lived at the farm where Russell's is today, but his store was located about half a mile away, at the intersection of Route 20 and Pelham Island Road, in Wayland Center. It stood near a grocery store called the Collins Market, along with the library, post office, and several churches. Everything was within walking distance. \"There were no cars in 1876, for convenience,\" I said. Tim held up a picture of the Russell's Provisions storefront for the camera to capture. We were filming a documentary about our family business, aiming to tell the story of how our 144-year-operation\u2014one of the oldest garden centers in the country\u2014tackled the challenges of the pandemic by changing our business dramatically. For us, the family history was a central motivation for maintaining the garden center through the initial closures in March 2020, when we experienced more than a million dollars in losses. We worried that we might have to close the business altogether. Genevieve asked, \"Was your grandfather a butcher too?\" \"Not at all\" I replied. I explained how my grandfather, Lewis Samuel Russell, was a farmer. Like his father, he grew vegetables and cut flowers on the family farm, and he also raised chickens and sold the eggs. In 1920, he opened Russell's Market in the space where we now sell garden tools\u2014right next to his house. At that point, cars were becoming more common, which meant that my grandfather could close the original location in town. It wasn't just my grandfather running the market, I explained. \"My Grammy, Ruth Russell, would add up customers' purchases on a little pad of paper and collect cash and make change out of her apron pocket.\" Genevieve asked me to pause for a moment and instructed the cameraman to zoom in on my face. She then asked, \"What was it like growing up on a farm?\" I described how I would visit my grandparents almost every day. I would play in the fields with my sisters and cousins, while my grandfather and great uncle worked nearby planting, weeding, and picking crops. At that point, my parents were involved with the business, so we would often stop to see them in the office, before heading to Grammy's yellow house, which still stands along Route 20. She'd give us fresh bread and sweets that she'd cooked on the old black coal stove. In the evenings, when my grandparents babysat for us, we'd watch Lawrence Welk and Carol Burnett on the television as they counted the cash from the day at their kitchen table. Family and business were inseparable. \"They'd hide the cash in an oatmeal box in the cupboard,\" I said. \"Once it was full, my grandma would put it in her bra and ride the bus to deposit it in the bank.\" Tim flipped the page of an album from 1965 to reveal a picture of my dad, Lewis Samuel Russell Jr., watering rows of flowers growing in our greenhouses. The cameraman zoomed in with his lens. My dad joined the business after he returned from the Korean War. By then, a significant part of the business revolved around wholesaling Five Generations of Russell's Garden Center Elizabeth Russell-Skehan Facing page: Russell's Garden Center has been a family-owned fixture in Wayland, Massachusetts, for five generations. PHOTOS COURTESY THE AUTHOR; USGS MAP FROM HARVARD UNIVERSITY, HARVARD MAP COLLECTION RUSSELL-SKEHAN, E. 2021. FIVE GENERATIONS OF RUSSELL'S GARDEN CENTER. ARNOLDIA, 78(5-6): 10-12 cut flowers to florists in the Boston area. My mom, Charlotte, worked as a bookkeeper and also managed the flower deliveries. Twice a week, she would load my sisters and me into the van and deliver flowers. We loved helping her carry the bunches of fresh flowers into the stores. After the energy crisis of the 1970s, we stopped growing cut flowers and closed our greenhouses every winter to conserve heat and save money. With specialization, airplanes and trucks could bring cut flowers from the southern regions of the United States and overseas, so Russell's stopped selling wholesale. My uncle had built several greenhouses, and my dad recommissioned them for growing annuals and vegetables. This transition was the start of the garden center as we know it today\u2014 and was yet another instance of the business evolving in response to changes in the market and technology. \"Because we were located on Route 20, we had plenty of customers driving by to stop in,\" I told the camera. \"We added houseplants, cactus, poinsettias, and potted mums and began selling more Christmas trees, wreaths, and fresh floral arrangements.\" At that point, my dad hired his best friend, Hugh McKenzie, who started the Garden Shop. Hugh added tools, fertilizers, and insecticides, along with garden statuary and supplies for birds. My mom worked long hours, too, and expanded the offerings to include vases, pots, silk flowers, candles, Christmas ornaments, and d\u00e9cor. At noon, Genevieve suggested we take a break. During the interview, her plan for structuring the film had shifted, and she wanted to run the idea past me. \"Mom,\" she said, \"I've decided to start with the history of Russell's before we go into the story of everything you all did to overcome the pandemic.\" I agreed that this was a great idea. We had already decided that the last thirty minutes of our movie would be about the remarkable response from our community once we were able to reopen the business in the spring of 2020, after more than a month of closure. We found that the community embraced gardening with newfound enthusiasm\u2014and in the end, Russell's not only survived 2020 but thrived. With the camera rolling again, Genevieve asked when Tim and I joined the company. Tim told the story of us joining in 1986. \"I'm a recovering mechanical engineer,\" he joked, \"and Elizabeth's expertise is in marketing and advertising. I quickly learned that this was a lot more fun than sitting in an office all day.\" I explained how, at this point, I'm delighted that our son, Dan Skehan, has joined us full time. He is the fifth generation to work at Russell's. With a background in accounting, human resources, and financial management, he was instrumental in helping us figure out how to stay in business through 2020. He secured payroll protection loans and helped us furlough and then rehire and train our employees. Moreover, he kept abreast with ever-changing guidelines from the Center for Disease Control and the State of Massachusetts. \"He remained calm and added a wealth of knowledge,\" I explained. \"I'm not sure we'd still be in business if we didn't know that Dan would be here to continue the legacy of Russell's Garden Center.\" Elizabeth Russell-Skehan is the president and vice president of marketing at Russell's Garden Center. They are now editing a full-length feature documentary film called Growing Through Covid- 19. To watch a trailer or to donate to the film, visit www.growingthroughcovid19.com. 12 Arnoldia 78\/5-6 \u2022 October 2021 STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION (Required by 39 U.S.C. 3685) 1. Title: Arnoldia. 2. Publication number: 0004-2633. 3. Filing date: September 22, 2021. 4. Frequency: Quarterly. 5. Number of issues published annually: 4. 6. Annual subscription price: $20.00 domestic; $25.00 foreign. 7-8. Address of offices of publication, publisher, and editor: The Arnold Arboretum of Harvard University, 125 Arborway, Boston, Suffolk County, MA 02130-3500. 9. Full names of publisher, editor, and managing editor: The Arnold Arboretum of Harvard University, publisher; Jonathan Damery, editor. 10. Owner: The Arnold Arboretum of Harvard University. 11. Known bondholders, mortgagees, and other security holders owning or holding 1 percent or more of total: None. 12. The purpose, function, and nonprofit status of this organization and the exempt status for federal income tax purposes have not changed during the preceding 12 months. 13. Publication name: Arnoldia. 14. Issue date for circulation data below: June 29, 2019. 15. Extent and nature of circulation. a. Average number copies each issue during preceding 12 months: 1,720. Actual number copies of single issue published nearest to filing date: 1,750. b. Paid and\/or requested circulation: (1) Paid outside-county subscriptions: Average: 28. Actual nearest to filing date: 25. (2) Paid in-county subscriptions. Average: 1,181. Actual nearest to filing date: 1,193. (3) Sales through dealers and carriers, street vendors, and counter sales: None. (4) Other classes mailed through the USPS: None. c. Total paid and\/or requested circulation. Average: 1,209. Actual nearest to filing date: 1,218. d. (1)(2) (3) Free distribution by mail. Average: 191. Actual nearest to filing date: 196. (4) Free distribution outside the mail: Average: 250. Actual nearest to filing date: 250. e. Total free distribution: Average: 441. Actual nearest to filing date: 446. f. Total distribution: Average: 1,650. Actual nearest to filing date: 1,664. g. Copies not distributed. Average: 70. Actual nearest to filing date: 86. h. Total. Average: 1,720. Actual nearest to filing date: 1,750. i. Percent paid and\/or requested circulation. Average: 73%. Actual nearest to filing date: 73%. I certify that all information furnished on this form is true and complete. Jonathan Damery, Editor."},{"has_event_date":0,"type":"arnoldia","title":"The Resilient Trees of Flower City","article_sequence":5,"start_page":13,"end_page":15,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25740","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160a76c.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Quinn, Mark","article_content":"On March 4, 1991, I awoke to a knocking on my door. A coworker from the Monroe County Parks Department in Rochester, New York, planned to pick me up early to go to a trade show in Syracuse. When I glanced at the clock, however, I realized the power was out. The clockface was blank. I dressed quickly in the dark, and when I stepped out the front door, I found that the day's agenda was completely different than planned. My coworker had indeed arrived to pick me up, but looking down the street, I saw that ice covered everything. My twenty-five-foot-tall white birch (Betula papyrifera) was bent over, with the tip touching the ground. (This tree later sprang back, showing the amazing resilience of trees to crises.) We headed for Highland Park, the historic arboretum on the south side of Rochester, where we both worked as horticulturists. After multiple turnarounds due to trees blocking the road, we finally arrived at the Highland Park production greenhouses. The scene that met us was shocking. A huge limb from a one-hundred-year-old European beech (Fagus sylvatica) had fallen on our turn-of-the-century glass greenhouse. Like most of the largest trees in the park, this beech dated to the early 1890s and was planted by horticulturist John Dunbar according to plans drafted by Frederick Law Olmsted. We immediately set to work removing the limb and closing the hole in the damaged greenhouse, stapling poly film to the cypress bars in an attempt to save the delicate orchids inside. As we worked to keep the plants from freezing, we could hear the occasional snap of limbs breaking elsewhere in the park, but we still had not fully comprehended the scale of devastation around us. Rochester has a special affinity for trees. In the early 1800s, it was dubbed the Flour City, as waterpower of the Genesee River was used to grind enormous amounts of flour that was then shipped via the Erie Canal. By the second half of the century, however, Rochester became the Flower City, home to many of the country's largest and most prosperous nurseries. Two nurserymen played an especially pivotal role: George Ellwanger and Patrick Barry, owners of the successful Mount Hope Nursery, which they established around 1840. In 1888, Ellwanger and Barry donated land from their nursery grounds to the city to be used as a public park. Later named Highland Park, this land occupied a highpoint overlooking the city and the southern tier hills. Olmsted was enlisted to design a system of parks for Rochester, including North Park (now Seneca Park) and South Park (now Genesee Valley Park). Considering the interest that local nursery owners had invested in tree cultivation, Olmsted designed Highland Park as an arboretum. Many of the specimens to be planted were donated by Ellwanger and Barry. Park Superintendent Calvin Laney began acquiring additional plants for the park, but it soon became clear that more horticultural help was required. Dunbar was hired in 1891 to oversee the plant collections in the park. He quickly forged relationships with other prominent horticulturists, including Charles Sprague Sargent of the Arnold Arboretum. The similarities between Highland and the Arnold are not just superficial. Both arboreta were designed by Olmsted and were envisioned as features within larger park systems. Both have the distinct feel of an Olmsted design, with curving paths following the contours of the landscape. Dunbar and the horticulturists who followed him maintained an active relationship with Sargent and others at the Arnold. For many decades, The Resilient Trees of Flower City Mark Quinn QUINN, M. 2021. THE RESILIENT TREES OF FLOWER CITY. ARNOLDIA, 78(5-6): 13-15 LIBRARY OF CONGRESS, GEOGRAPHY AND MAP DIVISION Highland Park 15 the institutions exchanged plant material, supporting research at both sites. As time passed, the products of these efforts matured into beautiful collections. In Rochester, the public has come to expect these large, well-maintained trees throughout our arboretum and park system. Still, as a community of tree lovers, we often take for granted the tremendous asset left by our predecessors\u2014until crisis strikes. The ice storm of 1991 was one of these events. Having saved the orchids, staff turned their attention to assessing the damage to the arboretum. It seemed that almost everything in the collection was either damaged or destroyed. At first, opening roads and paths so people could get around was the priority. This effort to restore access took days. As the work progressed, we started to look at individual specimens and, to our dismay, found many of our most celebrated trees were no more. One public favorite, a katsura tree (Cercidiphyllum japonicum), looked like the last few feet of every branch was broken and hanging. The tree had been received in 1919 from the nursery of Leon Chenault, in Orleans, France. Once the forestry team addressed safety issues elsewhere in the landscape, they turned to the katsura, spending days expertly trimming off every broken limb. Today, three decades later, no evidence of the trauma remains. The katsura has returned bigger and better than ever. The saddest loss for me was a Persian ironwood (Parrotia persica), which had been received from Veitch Nursery, in England, in 1892. The specimen\u2014perhaps my favorite tree in the park\u2014was fascinating, forming an impenetrable maze of eight- to sixteen-inch trunks with gray-green mottled bark. It had been completely uprooted and was lying on the ground. I remember cutting up the branches and wondering if another specimen as impressive as this one existed anywhere. Yet, sometimes having too much to do can play in our favor: with thousands of trees down and in need of work, our team deferred grinding stumps until later. That spring, dozens of new shoots sprouted from the overturned Parrotia stump. Over time, our team thinned the shoots, allowing space for some to grow. Now thirty years have passed, and the plant is once again a tangle of trunks\u2014 again one of my favorites. While so many trees were damaged and lost, others weathered the storm with remarkable ease. Walking through the park, you come to an impressive pair of zelkovas (Zelkova serrata), found in the valley behind the historic Lamberton Conservatory. One of the trees was received in 1899 from Thomas Meehan & Sons, in Germantown, Pennsylvania, and the other arrived in 1919 from the Arnold Arboretum. These trees stood strong against the ice. Likewise, at the corner of Highland Avenue and Goodman Street, a dawn redwood (Metasequoia glyptostroboides) did the same. The tree was grown from seed distributed by the Arnold Arboretum in 1948, when this newly identified species was first introduced to North America. The dawn redwood flexed under the weight of the ice but bounced back with little damage. Despite the losses to the ice storm, Highland Park recovered. Every morning, I drive through the pinetum, which includes hundreds of varieties of mature evergreens\u2014an uncommon and, I think, underappreciated asset for a city park. The pinetum is particularly impressive in the winter with snow on the trees, giving the impression of being in an evergreen forest far north of Rochester. As I pull into my parking spot, I glance to a nearby hill where I see two magnificent fernleaf beech trees (Fagus sylvatica 'Asplenifolia') standing amongst a grouping of beech trees of other varieties. These two were donated from Ellwanger and Barry's Mount Hope Nursery in 1892. Looking to the left, I can see an American chestnut (Castanea dentata), about thirty feet tall and starting to succumb to blight, a remnant of a former crisis. Each of the trees stands as a living history\u2014a testament not only to their own resilience but to the commitment of the generations of horticulturists who have built and stewarded the plant collections in Flower City. Mark Quinn is the superintendent of horticulture for Monroe County Parks, in Rochester, New York. He oversees the cultivation and care of the botanical collection at Highland Park and all the parks throughout the County Parks System. Facing page: The author stands with one of the celebrated trees at Highland Park\u2014a katsura tree (Cercidiphyllum japonicum) received in 1919. PHOTO COURTESY THE AUTHOR"},{"has_event_date":0,"type":"arnoldia","title":"The Prince Family: Pioneers of American Horticulture","article_sequence":6,"start_page":16,"end_page":23,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25741","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160ab6f.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Casscles, J. Stephen","article_content":"It was a beautiful day on August 1, 1782, when Prince William Henry, the third son of King George III, was received at the home and gardens of William Prince Sr. in Flushing Landing, New York. The American Revolutionary War had effectively ended the year before when the British surrendered at the Battle of Yorktown. Yet, the sixteen-year-old visitor, who would, in 1830, rise to the throne as King William IV, had come to present a stand of colors to the King's American Dragoons, encamped three miles to the east of the Princes. The British soldiers were invited for a barbecue of a whole roasted ox at the Prince home, not the kind of warm reception that an American patriot would have given to a future British monarch and his troops. Prince was a nursery owner, almost forty years older than William, and the visit suggests the prominence of both Prince and the nursery. During the visit, Prince and William discussed their shared interest in growing and breeding plums, a specialty of the nursery. Plums were a critical fruit crop because they could be dried and stored for long periods and used as a nutritious food by the British Navy. Prince had introduced new plum varieties to Long Island, observing the acclimatization of the green gage plum (a common form of Prunus domestica). He even developed new varieties of plums, including 'Yellow Gage', which he would officially introduce the year after William's visit. In 1789, another group of illustrious visitors stopped at Prince's nursery: the newly elected president of the United States, George Washington, and his entourage of vice president John Adams, New York governor George Clinton, and the president of the Continental Congress, John Jay. Washington was less impressed with the nursery than William had been. He noted a large number of young fruit trees but described the shrubs as \"trifling\" and the flowers as \"not numerous.\" Flushing had been under British military occupation for the past seven years, and little plant material could be shipped during those long years of hostility. Nonetheless, by the 1790s, the Prince Nursery was likely the largest propagator of grafted fruit trees in the United States. It would grow to become even more: a center of horticultural learning. The Prince family's horticultural enterprise originated with William Prince's father, Robert, who was born in the 1690s. (His birth year has been variously presented as 1692 and 1699.) By 1723, Robert had begun collecting, growing, and propagating trees for his fruit farm. The plants included varieties of apples, pears, plums, nectarines, peaches, cherries, and small fruits. Throughout Robert's life, the nursery slowly evolved into a vibrant commercial operation, occupying eight acres directly south of what is now Northern Boulevard. This first Prince homestead was a beautiful structure with rounded shingles, set in a bank of flowering shrubs on the western edge of his property, next to the Flushing Creek. Flushing\u2014in northern Queens County\u2014was an ideal location for a nursery that would grow to become national in scope. It sits on the Long Island Sound, where winters are milder than most other parts of the state and where summers are cooler and less humid than colonial centers to the south. Flushing boasted high-quality topsoil, rich and fertile, with few stones. An underlying subsoil provided good water drainage while retaining sufficient moisture to allow plants to grow quickly. Flushing's location relative to the Port of New York meant that plants could readily be shipped to other parts of the country and Europe. Moreover, Flushing benefited from the cultural and financial rise of New York City. These factors would, in the nineteenth century, induce many other prominent nurseries to establish operations in Flushing. Robert and his wife, Mary Burgess, had six children. Their oldest son, William, took over the nursery by 1745, the year before Robert's death. Under William's leadership, the nursery ultimately expanded to twenty-four acres. The diversity of plants increased, as did the total sales. At the time, the standard American practice for propagating fruit trees, especially peaches (Prunus persica), was to grow seedlings and not to graft a tree to a suitable rootstock. Because of this seed-grown method, the quality of orchard trees was unknown until they came to maturity. Prince realized the commercial value of predictability and often budded or grafted his fruit trees to keep the variety true. The nursery expanded quickly between 1750 and the beginning of the American Revolutionary War in 1776. William published his firstknown notice of advertisement on September 21, 1767, which stated, \"For sale at William Prince's nursery, Flushing, a great variety of fruit trees, such as apple, plum, peach, nectarine, cherry, apricot and pear. They may be put up so as to be sent to Europe. Capt. Jeremiah Mitchell and Daniel Clements go to New York in passage boats Tuesdays and Fridays.\" The nursery's first-known catalogue appeared in 1771, a single-page broadsheet. The list contained over 230 plant selections, which was sizable for a nursery in colonial America. In addition to fruit crops, the offerings included evergreen trees, timber trees, and shrubs. Among the ornamental selections, tulip trees (Liriodendron tulipifera) and lilacs (three varieties, presumably Syringa vulgaris) were among the most expensive. An advertisement published in the New York Mercury, dated March 14, 1774, stated that William Prince was selling more than one hundred Carolina magnolias (Magnolia grandiflora) that were over four feet tall, raised from seed. He also advertised ninefoot- tall catalpas (Catalpa speciosa). The Revolutionary War halted the shipment of Prince's plants to most parts of the American colonies, except for areas under British control, such as Manhattan, Brooklyn, Long Island, and parts of the South. These wartime closures hurt the business. Reports variously state that somewhere between three thousand to thirty thousand grafted cherry trees were either purchased or confiscated by the British, to be used as hoops for making barrels. Yet, the Princes were likely British Loyalists and benefited from military protection. In fact, William's daughter Sarah married a British Army Major, Charles McNeill, who resigned from his military service after the war. And the British General Lord Howe ordered army units to guard the nursery, posting soldiers at the entrances. When George Washington visited the Princes with his entourage in 1789, his assessment of the poor quality and low diversity of the ornamental plants may suggest that nursery was still recovering from the war. Yet, by the summer of 1791, secretary of state Thomas Jefferson and his fellow Democratic-Republican James Madison of Virginia visited the nursery and reported more favorably. The men were touring New York and New England to study botanical curiosities, wildlife, and historic battlefields. They maintained that the tour was for health reasons and scientific exploration. Yet, those versed in politics noted that the trip was conducted through the country's Federalists strongholds of New York and New England instead of areas dominated by Jefferson's political base of Democratic-Republican support. Jefferson desired to improve domestic agriculture and arranged the nursery stop to discuss his ideas with William. Among the topics, they talked about Jefferson's vision for promoting the cultivation of sugar maples (Acer saccharum) for syrup production. Jefferson also took the opportunity to order plants for himself: sugar maples, highbush cranberries (Viburnum trilobum), balsam poplars (Populus balsamifera), and Beurre Gris pears (a variety of Pyrus communis). Later, he expanded his order to include stone fruits and nut trees, along with an array of ornamental trees, shrubs, and roses. As the United States grew towards the close of the century, so did the Prince Nursery. By 1793, William Prince, at the age of sixty-eight, turned over operations to his sons Benjamin and William Jr. Benjamin maintained the original family nursery for many years, calling it the Old American Nursery, but it was William Jr. who became the primary mover of the family business in the third generation. In 1793, he purchased twenty-four acres directly northeast of the original nursery. There, on the banks of \u222b Flushing Creek, he established his Linnaean Botanic Garden and Nursery. He designed it as a showplace to educate the public on botanical matters, including native plants, new varieties bred in the United States, and plants imported from Europe and farther afield. William Jr. and his son William Robert Prince took up the cause of identifying and describing plant material so that it could be offered to the public\u2014and they were highly invested in acquiring newly introduced species. In 1804, for instance, Meriwether Lewis and William Clark embarked upon the Missouri River to explore the recently acquired Louisiana Purchase. The expedition had been commissioned at Jefferson's request, and when the explorers returned east, they came bearing seeds and other botanical collections. The Princes were among the first nursery operators to grow and distribute plants from the expedition, and the Oregon grape holly (Mahonia aquifolium) became one of their most successful new products. The Princes were also among the first American nurseries to offer ornamental species from East Asia, like the golden rain tree (Koelreuteria paniculata), lacebark elm (Ulmus parvifolia), and Chinese wisteria (Wisteria sinensis). By the mid-1830s, William Jr. had ten nursery outbuildings, of which several were greenhouses that contained tropical and subtropical plants from Africa and Asia. Visitors could pay an admission fee to experience the warmth and humidity of the greenhouse\u2014a rewarding respite to escape the dark, cold New York winter. The nursery catalogue listed ten tropical hibiscuses (Hibiscus) and two gardenias (Gardenia) that bloomed in their greenhouses. Prince grew tropical fruits and flowers specifically for winter viewing. For variety, they also exhibited insectivorous plants such as sundew (Drosera), pitcher plant (Sarracenia), and Venus flytrap (Dionaea). Moreover, in 1833, The New-York Annual Register reported that the gardens and nursery covered up to forty In 1793, William Prince Jr. purchased twenty-four acres alongside the original nursery, naming the new property the Linnaean Botanic Garden and Nursery. In the decades to come, a cohort of nurseries would open in Flushing, including Parsons Nursery and Bloodgood Nursery, both mapped nearby in 1841. SMITH, 1841\/LIBRARY OF CONGRESS, GEOGRAPHY AND MAP DIVISION ies cultivated in America, other than apples. (While the father and son intended to treat apple cultivation with a third volume, the work was never published.) Like A Short Treatise on Horticulture, this book was widely read in America and became influential among aspiring horticulturalists. Moreover, the Princes paid particular attention to the nomenclature of the fruits covered in all of the publications, untangling confusion occurring in the field. This interest in the accurate classification of horticultural plants began with the work of William Sr., and it was among the family's most significant contributions to American horticulture. As a testament to William Jr.'s interest in classification, he displayed in his home a bust of Carl Linnaeus, the Swedish botanist who formalized the modern system of botanical nomenclature. William Jr. received the statue in a presentation by New York governor DeWitt Clinton at a meeting of European and American scientists to honor Linnaeus's birthday in 1823. A simultaneous celebration in Virginia was officiated by Thomas Jefferson, an honorary member of the Linnaean Society of Paris. By the time William Jr. died in 1842, Flushing had become a vibrant center for American horticulture. Bloodgood Nursery had been established there in 1798 and would become known as a specialist in maples. (A common Japanese maple even bears the name of the nursery: Acer palmatum 'Bloodgood'.) G. R. Garretson Nursery, a seed company specializing in flowers and vegetables, was established in 1836 and would grow to cover one hundred acres, supplying wholesale seeds to nurseries across the United States and offering retail via mail order. But the most famous of these newer operations was Parsons Nursery, established in 1838; the Parsons family would later play a central role in introducing plants from East Asia, especially Japan. Meanwhile, William Robert had been assuming increasing responsibility for the Linnaean Botanic Garden and Nurseries. In the 1820s, he expanded the nursery, purchasing three large parcels so that his land holdings may have totaled up to 113 acres. These properties were located adjacent to a house he bought for himself in 1827. The home had a wide center hall, \u222b 20 Arnoldia 78\/5-6 \u2022 October 2021 acres and contained approximately ten thousand species of trees and plants, with particular attention devoted to grapes and mulberry trees. Visitors had free access to the outdoor gardens every day, except for Sundays. At the same time, the commercial operations of the nursery expanded rapidly, as evidenced by William Jr.'s increasingly thicker plant catalogues. He also began to subdivide the products among smaller specialized catalogues. In addition to his standard Annual Catalogue for Fruit and Ornamental Trees and Plants, which covered his earlier offerings, he began to issue catalogues that focused on items such as bulbous flowers and tubers, greenhouse plants, chrysanthemums, and vegetable and flower seeds. William Jr. attracted additional attention in 1828 when he published one of the first strictly horticultural books to come from the United States: A Short Treatise on Horticulture: Embracing Descriptions of a Great Variety of Fruit and Ornamental Trees and Shrubs, Grape Vines, Bulbous Flowers, Green-House Trees and Plants, &c. The book described all the plant offerings at the Linnaean Botanic Garden and Nursery, in some sense serving as an extended advertisement. The treatise also comprehensively covered horticultural topics, such as planting, pruning, and propagation. It even included information about soil preferences and methods for fungal disease control. Over the next three years, William Jr. worked with his son, William Robert, on two additional books, for which his son was the primary author. The first, A Treatise on the Vine, was published in 1830 and was the first significant book written in America on grape cultivation. The Princes had systematically tested scores of European grape varieties (Vitis vinifera), along with improved varieties of native North American grapes (like V. labrusca and V. riparia), and interspecific hybrids. The book described over two hundred European grape varieties and eighty American. This work helped to establish viticulture as a fullfledged branch of American horticulture, and for William Robert, grape breeding and cultivation remained a lifelong interest. The second book, The Pomological Manual, published in 1831, was a two-volume cyclopedia that attempted to catalogue all fruit varietwith two solid Dutch doors on either end and a bust of Linnaeus (likely from his father) on a bracket against the wall. The house's formal gardens contained two ginkgos (Ginkgo biloba), which were among the oldest in the country, and an old cedar of Lebanon (Cedrus libani) that the Princes had imported from France. Under William Robert's leadership, however, the business began to struggle. In the 1830s, he speculated heavily in the domestic silk industry and may have been a key contributor to the skyrocketing prices for mulberry trees (Morus alba), the food source for silkworms. He imported more than one million mulberry trees from France in 1839, and shortly afterward, the price for mulberry trees crashed. When this venture failed, the Princes could not keep up with mortgage payments on the nursery, and by 1841, they lost the Linnaean Botanic Garden and Nurseries in foreclosure. These events spawned a bitter controversy with the property's new owner, Gabriel Winter, who was married to one of William Jr.'s cousins. Although William Robert continued to raise and sell plants from an adjacent nursery property, he and Winter competed in horticultural publications over the right to sell plants as the Linnaean Botanic Garden and Nurseries. Ultimately, the Princes kept the name, and Winter sold the remaining plant inventory and subdivided the original property for housing development. By 1846, the finances at the new Prince nursery began to stabilize, and William Robert published Prince's Manual of Roses, his third and final significant contribution to horticultural literature. At his new botanic garden, William Robert grew over seven hundred rose varieties, and the book provided detailed descriptions of varieties and featured many roses from China. He also included information about horticultural care and propagation. It was one of the very best works on this subject. Still, it was eclipsed in popularity by Samuel B. Parsons's book published the following year: The Rose: Its History, Poetry, Culture, and Classification. Parsons\u2014the proprietor of Parsons Nursery in Flushing\u2014ultimately revised his book as Parsons on the Rose: A Treatise on the Propagation, Culture, and History of the Rose. The competition between these books suggests the horticultural foment that was occurring in Flushing during this period. William Prince Jr. and his son William Robert Prince (above) authored seminal American horticultural manuals. In A Treatise on the Vine, published in 1830, they promoted new grape varieties, including 'Isabella', which became a favorite of American viticulturists. HEDRICK, 1908 AND 1911\/ARCHIVE OF THE ARNOLD ARBORETUM 22 Arnoldia 78\/5-6 \u2022 October 2021 Later, William Robert went on two extended botanical expeditions, to California (in 1849) and Mexico (in 1850). While these trips suggest that the business was doing reasonably well, William Robert began to gradually withdraw from the day-to-day management of the nursery around 1855, at the age of sixty. Instead, he devoted his energy to other botanical interests, including research on botanical medicinal remedies. He also continued to breed and evaluate new varieties of fruits and ornamental plants, especially grapes, strawberries, and roses. His oldest son, William III, meanwhile assumed increasing responsibility for the enterprise. William Robert's career reflected the changes that were going on in the American horticultural community. His father had been a founding member of the New York Horticultural Society in 1818 and joined the Massachusetts Horticultural Society after it was established in 1829, but he was also a member of the Linnaean Society of Paris, the Horticultural Society of London and Paris, and the Academy of Georgofili, based in Florence, Italy. William Robert invested his energy into the increasingly sophisticated American horticultural societies rather than those in Europe. He contributed many articles to the leading American agricultural magazines of the day, such as The Rural New Yorker and Gardener's Monthly. Moreover, he was a member of the American Institute of the City of New York and the American Pomological Society. On March 28, 1869, William Robert died at his home in Flushing, and as it turned out, the esteemed business died with him. William III had enlisted for the United States Army during the Civil War, and he chose to remain in the military. William Robert's second son, LeBaron Bradford, pursued a career in law and politics. Gardener's Monthly printed a two-page obituary for William Robert. It was a sad and respectful tribute to his horticultural brilliance while nonetheless remarking on his combative personality. Meanwhile, the Massachusetts Horticultural Society issued a full resolution commemorating his life as a \"pioneer in the field of horticulture,\" a title that seems equally appropriate for the three generations of Princes that came before him. In 1939, efforts were made to move William Robert's house to the site of the New York World's Fair, to demonstrate a historic colonial homestead, but the campaign came to no avail. Later, New York City park commissioner Robert Moses rejected a proposal to move the structure to Flushing Meadow Park. Moses's vision for a \"modern city\" had little space for old wooden buildings. In its last few years of use, the structure served as a rooming house and a club. The shabby, unpainted building was then boarded up and surrounded by billboards and a gas station. The house was torn down in 1942. Of course, by that point, the lush greenhouses that once welcomed winter visitors had long ago disappeared, and the nursery property had been subdivided and sold for development. Yet, the 150-year story of the Prince family lives on today. The family built a foundation for commercial horticulture in the United States. They championed the cultivation of plants from across the country and around the world, and their publications promoted best practices in horticulture. They even helped with establishing a more systematic approach for horticultural nomenclature. Moreover, the success of the Prince nurseries is inextricably linked to the subsequent generation of horticulturists who established businesses in Flushing. This expanding group of nursery owners became leaders in their own right. In this way, a horticultural legacy that began with one family who lived on the edge of Flushing Creek became a national and international story. Acknowledgment I'm grateful for the support of Susan Lacerte, who recently retired as executive director at the Queens Botanical Garden, located near the former Prince Nurseries. Susan's knowledge of horticulture in Flushing, both present and historical, has been an inspiration. References Cornett, P. 2004, January. Encounters with America's premier nursery and botanic garden. Twinleaf: 1-12. Downing, A. J. 1845. The fruits and fruit trees of America: Or the culture, propagation and management in the garden and orchard of fruit trees generally; with descriptions of all the finest varieties of fruit, native and foreign in this country. New York: Wiley and Putnam. \u222b The Prince Family 23 Gager, C. S. 1912, October. The first botanic garden on Long Island. Brooklyn Botanic Garden Record, 1(4): 97-99. Hedrick, U. P. 1911. The plums of New York (Report of the New York Agricultural Experiment Station). Albany, N.Y.: J. B. Lyon Company. Hedrick, U. P. 1908. The grapes of New York (Report of the New York Agricultural Experiment Station). Albany, N.Y.: J. B. Lyon Company. Hedrick, U. P. 1925. The small fruits of New York (Report of the New York Agricultural Experiment Station). Albany, N.Y.: J. B. Lyon Company. Hedrick, U.P. 1933. A history of agriculture in the State of New York. Printed for the New York State Agricultural Society, Albany, N.Y.: J. B. Lyon Company. Hotchkiss, T. W. 1934. Prince house, Lawrence Street & Northern Boulevard, photographs, written historical and descriptive data. Dist. No. 4, Southern New York State, Historic American Building Survey, HABS No. 4-19. Jacobsen, A. and Williams, J. D. 2009. Prince family nurseries (ca. 1737- post- 1851). Bulletin of the Hunt Institute of Botanical Documentation, 21(1): 4-7. Johnson, J. 1887. The village of Flushing, map of desirable building lots, Flushing: A historical sketch. New York: John P. Stock, Printer. Manks, D. S. 1967. How the American nursery trade began. Plants & Gardens, 23(2). McGourty, F. 1967. Long Island's famous nurseries. Plants & Gardens, 23(3). Munsell, W. W. 1882. History of Queens County, New York, with illustrations, portraits, & sketches of prominent families and individuals. New York: Press of George MacNamara. Prince, B. and Mills, S. F. 1823. A treatise and catalogue of fruit and ornamental trees, shrubs, &c., cultivated at the Old American Nursery. New York: Wm. Grattan. Prince, W. 1771. To be sold, by William Prince, at Flushing-Landing, on Long-Island, near New- York, a large collection of fruit trees, as follows. New York: H. Gaine. Prince, W. 1790. To be sold, by William Prince, at Flushing-Landing, on Long-Island, near New- York, a large collection, as follow, of fruit trees and shrubs. New York: H. Gaine. Prince, W. 1825. Annual catalogue of fruit and ornamental trees and plants, bulbous flower roots, green-house plants, &c. &c., cultivated at the Linnaean Botanic Garden, William Prince, proprietor. New York: T. and J. Swords. Prince, W. 1828. A short treatise on horticulture: Embracing descriptions of a great variety of fruit and ornamental trees and shrubs, grape vines, bulbous flowers, green-house trees and plants. New York: T. and J. Swords. Prince, W. R. and Prince, W. 1830. A treatise on the vine; Embracing it history from the earliest ages to the present day, with descriptions of above two hundred foreign, and eighty American varieties, together with a complete dissertation of the established culture, and management of vineyards. New York: T. & J. Swords. Prince, W. R. and Prince, W. 1831. The pomological manual; or a treatise on fruits: containing descriptions of a great number of the most valuable varieties for the orchard and garden. New York: T. & J. Swords. Prince, W. R. 1846. Princes' Manual of roses: Comprising the most complete history of the rose, including every class, and all the most admirable varieties that have appeared in Europe and America, together with ample information on their culture and propagation. New York: Clark & Austen, Saxton & Miles, Wiley & Putnam, and Stanford & Swords. Ross, P. 1902. A history of Long Island: From its earliest settlement to the present time. New York: Lewis Publishing Co. Smith, E. A. and Hayward, G. 1841. The village of Flushing, Queens County, L.I.: nine miles east of the city of New York: lat. 40\u00b0 45' 1\"N, lon. 73\u00b0 09' 58\"W. [Flushing?: s.n., ?] [Map] Retrieved from the Library of Congress, https:\/\/www.loc. gov\/item\/2008620796 St. George's Episcopal Church, Baptismal Records, 1800- 1840, Flushing, N.Y., 135-32 38th Avenue, Flushing, N.Y., Rev. Wilfredo Benitez, Rector. Trebor, H. (Ed.) 1938, October. Garden center: The four Princes\u2014William of America. So This is Flushing. Flushing, N.Y.: Halleran. U.S. Department of Agriculture. 1976. The Prince family manuscript collection: A register of their papers, in the National Agricultural Library (Library list 101). Beltsville, MD: U.S. Department of Agriculture, National Agricultural Library. Waldron, R. K. 1958. Prince's plants. The Call Number, 20(1). J. Stephen Casscles is an attorney, winemaker, and horticultural writer living in the Hudson Valley. His publications include Grapes of the Hudson Valley and Other Cool Climate Regions of the United States and Canada, published by Flint Mine Press."},{"has_event_date":0,"type":"arnoldia","title":"Such a Fine Assemblage: The Jesup Collection of North American Woods","article_sequence":7,"start_page":24,"end_page":49,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25742","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160af27.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Mauz, Kathryn","article_content":"It was a beautiful day on August 1, 1782, when Prince William Henry, the third son of King George III, was received at the home and gardens of William Prince Sr. in Flushing Landing, New York. The American Revolutionary War had effectively ended the year before when the British surrendered at the Battle of Yorktown. Yet, the sixteen-year-old visitor, who would, in 1830, rise to the throne as King William IV, had come to present a stand of colors to the King's American Dragoons, encamped three miles to the east of the Princes. The British soldiers were invited for a barbecue of a whole roasted ox at the Prince home, not the kind of warm reception that an American patriot would have given to a future British monarch and his troops. Prince was a nursery owner, almost forty years older than William, and the visit suggests the prominence of both Prince and the nursery. During the visit, Prince and William discussed their shared interest in growing and breeding plums, a specialty of the nursery. Plums were a critical fruit crop because they could be dried and stored for long periods and used as a nutritious food by the British Navy. Prince had introduced new plum varieties to Long Island, observing the acclimatization of the green gage plum (a common form of Prunus domestica). He even developed new varieties of plums, including 'Yellow Gage', which he would officially introduce the year after William's visit. In 1789, another group of illustrious visitors stopped at Prince's nursery: the newly elected president of the United States, George Washington, and his entourage of vice president John Adams, New York governor George Clinton, and the president of the Continental Congress, John Jay. Washington was less impressed with the nursery than William had been. He noted a large number of young fruit trees but described the shrubs as \"trifling\" and the flowers as \"not numerous.\" Flushing had been under British military occupation for the past seven years, and little plant material could be shipped during those long years of hostility. Nonetheless, by the 1790s, the Prince Nursery was likely the largest propagator of grafted fruit trees in the United States. It would grow to become even more: a center of horticultural learning. The Prince family's horticultural enterprise originated with William Prince's father, Robert, who was born in the 1690s. (His birth year has been variously presented as 1692 and 1699.) By 1723, Robert had begun collecting, growing, and propagating trees for his fruit farm. The plants included varieties of apples, pears, plums, nectarines, peaches, cherries, and small fruits. Throughout Robert's life, the nursery slowly evolved into a vibrant commercial operation, occupying eight acres directly south of what is now Northern Boulevard. This first Prince homestead was a beautiful structure with rounded shingles, set in a bank of flowering shrubs on the western edge of his property, next to the Flushing Creek. Flushing\u2014in northern Queens County\u2014was an ideal location for a nursery that would grow to become national in scope. It sits on the Long Island Sound, where winters are milder than most other parts of the state and where summers are cooler and less humid than colonial centers to the south. Flushing boasted high-quality topsoil, rich and fertile, with few stones. An underlying subsoil provided good water drainage while retaining sufficient moisture to allow plants to grow quickly. Flushing's location relative to the Port of New York meant that plants could readily be shipped to other parts of the country and Europe. Moreover, Flushing benefited from the cultural and financial rise of New York City. These factors would, in the nineteenth century, induce many other prominent nurseries to establish operations in Flushing. Robert and his wife, Mary Burgess, had six children. Their oldest son, William, took over the nursery by 1745, the year before Robert's death. Under William's leadership, the nursery ultimately expanded to twenty-four acres. The diversity of plants increased, as did the total sales. At the time, the standard American practice for propagating fruit trees, especially peaches (Prunus persica), was to grow seedlings and not to graft a tree to a suitable rootstock. Because of this seed-grown method, the quality of orchard trees was unknown until they came to maturity. Prince realized the commercial value of predictability and often budded or grafted his fruit trees to keep the variety true. The nursery expanded quickly between 1750 and the beginning of the American Revolutionary War in 1776. William published his firstknown notice of advertisement on September 21, 1767, which stated, \"For sale at William Prince's nursery, Flushing, a great variety of fruit trees, such as apple, plum, peach, nectarine, cherry, apricot and pear. They may be put up so as to be sent to Europe. Capt. Jeremiah Mitchell and Daniel Clements go to New York in passage boats Tuesdays and Fridays.\" The nursery's first-known catalogue appeared in 1771, a single-page broadsheet. The list contained over 230 plant selections, which was sizable for a nursery in colonial America. In addition to fruit crops, the offerings included evergreen trees, timber trees, and shrubs. Among the ornamental selections, tulip trees (Liriodendron tulipifera) and lilacs (three varieties, presumably Syringa vulgaris) were among the most expensive. An advertisement published in the New York Mercury, dated March 14, 1774, stated that William Prince was selling more than one hundred Carolina magnolias (Magnolia grandiflora) that were over four feet tall, raised from seed. He also advertised ninefoot- tall catalpas (Catalpa speciosa). The Revolutionary War halted the shipment of Prince's plants to most parts of the American colonies, except for areas under British control, such as Manhattan, Brooklyn, Long Island, and parts of the South. These wartime closures hurt the business. Reports variously state that somewhere between three thousand to thirty thousand grafted cherry trees were either purchased or confiscated by the British, to be used as hoops for making barrels. Yet, the Princes were likely British Loyalists and benefited from military protection. In fact, William's daughter Sarah married a British Army Major, Charles McNeill, who resigned from his military service after the war. And the British General Lord Howe ordered army units to guard the nursery, posting soldiers at the entrances. When George Washington visited the Princes with his entourage in 1789, his assessment of the poor quality and low diversity of the ornamental plants may suggest that nursery was still recovering from the war. Yet, by the summer of 1791, secretary of state Thomas Jefferson and his fellow Democratic-Republican James Madison of Virginia visited the nursery and reported more favorably. The men were touring New York and New England to study botanical curiosities, wildlife, and historic battlefields. They maintained that the tour was for health reasons and scientific exploration. Yet, those versed in politics noted that the trip was conducted through the country's Federalists strongholds of New York and New England instead of areas dominated by Jefferson's political base of Democratic-Republican support. Jefferson desired to improve domestic agriculture and arranged the nursery stop to discuss his ideas with William. Among the topics, they talked about Jefferson's vision for promoting the cultivation of sugar maples (Acer saccharum) for syrup production. Jefferson also took the opportunity to order plants for himself: sugar maples, highbush cranberries (Viburnum trilobum), balsam poplars (Populus balsamifera), and Beurre Gris pears (a variety of Pyrus communis). Later, he expanded his order to include stone fruits and nut trees, along with an array of ornamental trees, shrubs, and roses. As the United States grew towards the close of the century, so did the Prince Nursery. By 1793, William Prince, at the age of sixty-eight, turned over operations to his sons Benjamin and William Jr. Benjamin maintained the original family nursery for many years, calling it the Old American Nursery, but it was William Jr. who became the primary mover of the family business in the third generation. In 1793, he purchased twenty-four acres directly northeast of the original nursery. There, on the banks of \u222b Flushing Creek, he established his Linnaean Botanic Garden and Nursery. He designed it as a showplace to educate the public on botanical matters, including native plants, new varieties bred in the United States, and pOn May 18, 1885, an important exhibition heralded as a \"noble gift to the city\"1 opened at the American Museum of Natural History in New York. Beneath the high ceilings of the exhibition hall, glass cases displayed 350 specimens as the Jesup Collection of North American Woods. Each was a whole log, about four and a half feet tall, still cloaked with bark as in life, with the upper half cut away to reveal the wood inside. Many of the specimens were accompanied by original watercolor illustrations of foliage, fruit, and flowers. A writer announced of the exhibit in Harper's Weekly, \"The average visitor will be impressed and surprised by the beauty of some and by the extreme oddity of others.\u2026 The various coloring of the woods, often rich and sometimes startling, and running into the most delicate shades, and the strength or grace or whimsicality of form, as traced in the divers[e] coursings of the grain, are matters to attract even the casual eye, and to stamp as absurd the hasty judgement which would say that a collection of logs can not be interesting.\"2 Over the coming years, the collection grew to include more than five hundred species. It represented the scientific and philanthropic vision of two noteworthy individuals: Morris Ketchum Jesup, one of the founders of the American Museum of Natural History, and Charles Sprague Sargent, the director of the Arnold Arboretum. The collection remained a cornerstone of the museum's exhibits for more than six decades. The fact that an exhibition of this magnitude could almost entirely vanish from the public memory seems almost improbable. Yet, the story of its exile is as intriguing as that of its origins. A Generous Friend On the occasion of the Philadelphia Centennial Exhibition of 1876, William H. Brewer, a professor of agricultural science at Yale University, observed, \"America has long been described by geographers and naturalists as the wooded continent, distinguished for the luxuriance and extent of its forests and the number of its arboreal species.\"3 At that time, scientists were beginning to comprehend the vastness of North American forests, but popular appreciation of this forest wealth lagged behind. At the Exhibition, audiences were introduced to displays of American woods and wood products through exhibits mounted by individual states and by the United States Department of Agriculture, which showcased specimens representing four hundred tree species from around the country.4 Such exhibits distilled an abstract general abundance into the remarkable variety of trees that comprised the country's forests. The Exhibition's millions of visitors vastly exceeded the number of people who had ever traveled across the country or explored its forested lands, and early efforts to organize around the idea of forest conservation took root at that gathering. At the time, there was not a museum in the country that possessed a similar, permanent exhibit that could perpetuate the transient awe from the Centennial Exhibition into an enduring educational mission. In 1880, such an exhibit\u2014but one even more monumental\u2014 became Jesup's vision for the American Museum of Natural History. A forest lover himself, Jesup was also keenly interested in the uses of forests and, increasingly, in the roles forests played in the wider landscape of human settlement and industry. Jesup and the museum's director, Such a Fine Assemblage: The Jesup Collection of North American Woods Kathryn Mauz Facing page: The Jesup Collection of North American Woods revealed the wonder and scientific diversity of North American forests by showcasing wood samples from more than five hundred tree species. As one commentator later said, it was \"a perfectly unique collection which cannot anywhere be repeated.\" AMNH RESEARCH LIBRARY DIGITAL SPECIAL COLLECTIONS, 31642 MAUZ, K. 2021. SUCH A FINE ASSEMBLAGE: THE JESUP COLLECTION OF NORTH AMERICAN WOODS. ARNOLDIA, 78(5-6): 24-49 26 Arnoldia 78\/5-6 \u2022 October 2021 Albert S. Bickmore, discussed the possibility of developing this exhibit at the museum for the expressed purpose of showcasing the contributions of American forests to industrial and artistic endeavors. In August 1880, while attending the annual meeting of the American Association for the Advancement of Science in Boston, Bickmore approached Harvard botany professor Asa Gray for advice. He described the museum's planned Department of Economic Botany, which was primarily to feature important products from the forests of the country. Gray directed him to interview Sargent, who at the time was in charge of the census of American forests for the Tenth Census of the United States. Bickmore spent an afternoon at Dwight House on Sargent's Holm Lea estate in the suburb of Brookline. Although Sargent was away conducting fieldwork, Bickmore toured the grounds and learned about the work Sargent was pursuing for the forest census. Bickmore soon wrote to Sargent in care of the Palace Hotel in San Francisco, where Sargent was briefly stopped along the last leg of his grand tour of western forests. As Bickmore explained, a \"generous friend\" of the museum wished to develop an \"instructive and attractive collection\" of the wood products of North American forests, \"placing it in a tangible, visual form before our citizens and our tide of visitors from all parts of the continent.\"5 Of course, that unspecified friend was Jesup, who would become the museum's president from 1881 until his death in 1908. His foresight had led him to Sargent, whose zeal and breadth of knowledge were positively suited to realizing this singular goal, and whose awareness of his own expertise prevented him from letting the opportunity pass to someone else. Jesup also sponsored other collections and many expeditions in varied fields of study during his tenure at the museum, and Sargent simultaneously expanded the Arnold Arboretum's living collection and pursued an astounding schedule of publication. Yet, the wood collection was seen as a crowning achievement during the lifetimes of both men. It was, according to one commentator, \"a perfectly unique collection which cannot anywhere be repeated.\"6 ARCHIVES OF THE ARNOLD ARBORETUM AMNH RESEARCH LIBRARY DIGITAL SPECIAL COLLECTIONS, 2A5200 The Jesup Collection emerged from the collaboration of Morris K. Jesup (right) and Charles Sprague Sargent. Jesup Collection 27 Unprecedented Activity Following his return from the west, Sargent met with Jesup and Bickmore in New York in the first week of November 1880. In response to the proposed project, he sent a seven-page letter describing his \"suggestions\" for the wood collection and its exhibition, which in effect were stipulations to guarantee his participation. Sargent believed that the collection should incorporate every tree species that grew naturally in the United States, even those that were of limited distribution or held little economic value. As a reflection of his recent and ongoing work on the forest census, he argued that only this approach would allow the collection's importance to be realized by both the public and scientists, who, he would later assert, \"will value it in proportion to its completeness.\"7 Further, Sargent insisted that the exhibit be arranged according to the botanical relationships of the species, following the organization of his report for the forest census, and that the labels should incorporate the data from his investigations as to each species' geographic distribution and the properties of its wood. He shared Jesup's interest in including foliage and fruit to illustrate the aspect of the living trees, as well as the products derived from the trees that were important to commerce and the trades.8 In essence, it would be a full-scale adjunct to his census report, one that Jesup hoped would also have popular appeal and that all concerned believed would be an asset to the museum.9 Sargent's primary role in the project was to direct and coordinate the field efforts and, later, to provide interpretation for the resulting specimens. By mid-December 1880, once a general plan for the collection was understood, he was becoming impatient to send collectors into the field.10 The first to be recruited were alumni of the forest census who were familiar with both the terrain and tree species they were to locate, as well as the rigors and routine of moving logs from the forests to the railroads for shipping. Some were in the field as early as January, and specimens began arriving at the museum in early March 1881. Charles Mohr, a physician and botanist who lived in Mobile, Alabama, was charged with finding trees in the Gulf Coast states. (Records show that the first specimen to be received may have been Yucca treculeana, or Spanish dagger, an arborescent species, if not precisely a tree, sent from Texas by Mohr.11) Samuel B. Buckley, a botanist and long-time resident near Austin, Texas, began collecting nearby and at points across the southern interior of the state. Allen H. Curtiss, a naturalist living in Jacksonville, Florida, was sent to explore southern Florida, the Florida Keys, and the interior Southeast; in his first season, Curtiss sent more than forty specimens, and he ultimately contributed more than any other collector. George W. Letterman, a schoolteacher and amateur botanist in Allenton, Missouri, began his work that spring in Arkansas, made numerous collections in southern and central Missouri, and later ventured as far as northeastern Texas and Louisiana. Henry W. Ravenel, an accomplished botanist of Aiken, South Carolina, sent specimens from the Piedmont and coast of South Carolina and Georgia that year. Starting in the fall of 1881, John H. Sears, a naturalist in Salem, Massachusetts, explored the \"Atlantic forests\" of northern New York state and eastern Massachusetts. For the first two years, Vermont botanist Cyrus G. Pringle traveled well beyond his home state to collect in Arizona, California, and the Pacific Northwest, and later sent logs of several species from Texas and northern Mexico, as well; second only to Curtiss in number of specimens sent, Pringle certainly traveled more extensively for the project than anyone else. The collecting corps came to include physicians, veterans of state geological surveys and departments of agriculture, itinerant botanists, horticulturists, foresters, several of Sargent's professional acquaintances in the lumbering and milling industries, Sargent himself, and even the collection's caretaker, Samuel D. Dill, at the museum. The majority of specimens were collected by a handful of men, but over time more than fifty individuals contributed material to the Jesup Collection. Sargent initially envisioned an ambitious schedule, entailing just one or two years to complete the explorations necessary to find and acquire the specimens.12 That, like the costs involved, turned out to be underestimated\u2014not 28 Arnoldia 78\/5-6 \u2022 October 2021 only were there unforeseen delays but more species in newly explored places were discovered over time, in part as a result of Sargent's own studies. As time went on, Jesup sometimes questioned the necessity for including extraneous, noneconomic species, noting to Sargent, \"Its completeness in a scientific or botanical sense, to my mind is secondary.\"13 To Bickmore privately, he observed that many tree species, \"while they may be rare and valuable in a scientific sense, are useless economically owing to the remote and inaccessible districts where they grow and the necessary cost of transportation to manufacturing centres.\"14 Sargent nonetheless continued to send collectors far afield and on special trips for newly discovered or rare species in the interest of amassing a comprehensive collection. He had taken on the project gratis, with an eye toward his own long-term interests in American forests. With the collection's scientific contributions as his priority, Sargent advised Jesup early in 1881, \"It is not too late for us both to retire altogether from the undertaking, which unless carried out largely will add neither reputation to the Museum, nor credit to the parties most interested.\"15 The project went on, and fifteen years later he emphasized the significance of the work to Jesup: \"The formation of your Collection, the publication of my book, and other causes have led to an unprecedented activity in dendrological exploration and study in all parts of the country and several new species of trees have been discovered.\"16 Sargent's aim was to represent the arboreal flora of the continent, and he wanted Jesup's vision to match his own. It Should Contain Every Tree As the sponsor of the collection, Jesup not only funded the collector's activities but organized logistics for travel and shipping. He was wealthy and generous, but disciplined and frugal in his philanthropy, interested to see that his money was well spent for the greatest benefit. To this end, he set as a goal keeping costs of travel and freight to a minimum, even zero, whenever possible. Nonetheless, the cost of transportation, shipping, and tracking the specimens across the country represented the majority of the project's expenses and occupied much of the correspondence between Sargent and the museum during these early years. In the early weeks of 1881, Jesup personally communicated with the officers of dozens of railroad and steamship companies in order to procure travel passes for the collectors and free shipping for the weighty specimens they were expecting to send to New York from points around the country. Because the favors granted were often specific to individual collectors, over certain routes, and good only for specified periods of time, this became for him a never-ending task that strained his ample reserves of tact and humility. Through Jesup's general success in securing waivers, Sargent could then assign collectors to regions where they could travel freely and ship at no or reduced cost. In practice, there were frequent misunderstandings on the part of station agents who were unaware of these unconventional arrangements or would not act on them. Specimens were sometimes shipped from points or by routes other than what had been agreed upon, exceeded the weights and dimensions originally anticipated, were delayed so long that they decayed in transit, or were occasionally even lost. The railroads, and Jesup, wanted definite parameters ahead of time, whereas Sargent better understood the idiosyncrasies and exigencies of field work and insisted that flexibility was necessary. It was Jesup's money, and indeed his reputation, at risk, and these overages and losses were routine points of contention between the two principals almost from the beginning.17 As the true scale of the task became apparent, Jesup questioned Sargent's early estimates about the cost of the project. He had initially thought that the collection could be completed for ten thousand dollars or possibly less,18 but that sum was exceeded before the end of the second year of work; total expenditures multiplied fivefold before the sixth field season and continued to grow from there.19 Although Sargent promised to proceed as economically as he could, he maintained his emphasis on the need for a complete and scientifically valuable set of specimens. Following one expensive expedition in 1885, for example, Sargent countered Jesup's objections, telling him, \"I hope you will not endeavor to separate practical value from Jesup Collection 29 scientific value in your mind when considering this collection. They cannot safely be separated. And it is because I have always refused to do this in the treatment of the matter that the collection is what it is, the best of its kind.\"20 Bickmore and Jesup at the museum recognized that ceding some control to Sargent (and absorbing additional expense) was necessary both to achieving that goal and to maintaining goodwill in general.21 Nearly two decades after the project's inception, as he and Jesup revisited this same familiar disagreement in 1899, Sargent argued, \"It should contain every tree described and illustrated in my Silva of North America.\"22 Although their differences in philosophy did not entirely fade over time, Jesup grudgingly found himself obligated to continue to subsidize these missions\u2014 well into the 1890s and, for a few species, even past the turn of the century\u2014rather than risk the appearance of incompleteness once so many others had been gathered. Early on he remarked to Sargent, \"To have our museum contain that which cannot be found at any other will fully compensate me for the cost.\"23 A Grand Showing Unlike the small blocks of wood Sargent prepared for his census investigations24 or the short logs cut lengthwise for display at the Centennial Exhibition, the museum's specimens were to be whole logs, over five feet long when collected, and of such diameters as necessary (from a few inches to three feet or more) to represent the best-grown examples of the trees. Collectors routinely shipped thousands of pounds of specimens at once, where certain individual logs could weigh hundreds of pounds when freshly cut. At the outset, Sargent anticipated that about four hundred species would need to be assembled, but that number increased by another one hundred or more over time. Within the year, Bickmore reported to Sargent, \"We have been frequently receiving the magnificent series of logs your agents have gathered until they make a grand showing in the cellar.\"25 After the first full year of fieldwork, nearly three hundred were in various states of preparation at the museum, with more arriving by the month.26 Incoming shipments were initially delivered to the museum's \"new building\" (opened in December 187727) on Manhattan Square, west of Central Park. When space became limited, the logs were directed instead to the historical Arsenal building, where the museum's collections were originally housed, near the eastern boundary of the park. When the logs were prepared in the field, collectors were careful to wrap each one in burlap or other \"bagging\" material, sometimes also in rawhide, and to construct crates in which the log could be shipped with ample padding to preserve the bark intact. Once at the museum's workshop, they underwent a lengthy process of preparation for eventual display. Because the logs were shipped \"green\" and were full of moisture, the primary concern was for drying them carefully to prevent \"checking\" or splitting that would ruin them for display. Bickmore himself devised a method of boring holes into the bottom of a log to allow the wood to \"season\" or dry out more evenly.28 Bickmore notified Sargent further, \"We have a fire under the boilers in the cellar constantly so that that is probably the driest room in the building, and the heat is gentle & slow and I believe particularly well adapted to preparing the fine logs that are now coming in, and I think there will be no necessity of having the specimens kiln dried, unless you have reason to suspect they contain destructive larvae.\"29 It was estimated that logs could lose up to half their weight in drying, and that thorough seasoning could sometimes require one or two years.30 Following the drying process, the logs were cut to a uniform fifty-six inches in height; the upper twenty-four inches was sawn longitudinally in half, and the top edge of the cut end was beveled, resulting in the grain of the wood being exposed in three directions. Finally, one half of the cut surface was finished with varnish to provide a clear view of the grain. Sargent requested that a diagram be made of each log to show the pattern of the bark, the widths of the sapwood and heartwood, and the growth rings apparent in cross-section;31 these data, as indicators of growth rate, were eventually reported for many species in Sargent's fourteen-volume Silva of North America, but the diagrams themselves have not survived. 30 Arnoldia 78\/5-6 \u2022 October 2021 Jesup's initial hopes that the collection would be ready for public viewing by the autumn of 1882 were not realized, but both he and Sargent agreed that the collection's \"value and permanence,\" from a scientific standpoint, and its \"beauty and usefulness\" to the public would be favored by postponing until all the specimens were fully seasoned, prepared, and labeled.32 The exhibit space dedicated to the Jesup Collection was intended to be on the third floor of the Arsenal, an area the museum regarded as \"dangerous\" even when exhibits had been open to the public there a decade earlier.33 Almost immediately, there were concerns about the combined weight of the specimens.34 When the walls of the building were observed to have to spread slightly by October 1882, the Department of Public Parks architect, Calvert Vaux, insisted that the excess weight be removed to comply with his specifications: not to exceed thirty-eight and a half tons, evenly distributed in the halls and the octagonal alcoves at each corner.35 At that time, there were 388 logs onsite and in preparation, with 60 more expected to \"complete\" the collection.36 This circumstance hinted at another persistent theme that would follow the collection through time: housing it would always present substantial, even prohibitive infrastructural challenges. Soon, the allotted hall at the Arsenal became a workshop and storeroom for the log specimens rather than their exhibit space. By the spring of 1883, construction at the museum's new building included the installation of \"a large glass case, in two sections, extending along the middle of the Lower Hall,\" meant to accommodate the log collection but necessarily displacing an exhibit of shells to another floor.37 By that autumn, there were two large cases, each 135 feet long, with six additional cases along the side.38 The initial delay of six months had extended to a full year, and even then, opening by the following year was in doubt. In February 1884, Sargent estimated that just 105 specimens were \"finished and ready\";39 in April, he wrote to Jesup and Bickmore to suggest delaying until the spring of 1885, when he thought that as many as 350 specimens would be fully prepared for exhibition.40 A Credit to the City With a date finally fixed for the exhibit's opening, Bickmore promoted it as \"the first effort yet made in this country to gather the native woods together in one collection on a scale commensurate with the extent of the new continent and the importance of its forests.\"41 Sargent had been at work on a condensed version of his census report, enumerating 412 species as The Woods of the United States, which would serve as a guidebook to the collection.42 In April, he reassured Jesup, \"The geographical labels will be finished this week. They have cost me an immense amount of labor & bother, but I think they will be a great success, and are certainly the best things of the kind ever attempted. I shall be in N.Y. next week, long enough to see that everything is properly arranged.\"43 In his annual report to the trustees of the museum, Jesup hoped that the collection \"will prove another popular attraction to the museum, and be the means of largely increasing the knowledge and information of the people on the subject of our forests, now demanding so large a share of public attention.\"44 The exhibit opened to visitors on May 18, 1885, to popular acclaim. In addition to 350 logs with their labels, the new exhibit featured about eighty watercolor illustrations of the foliage, flowers, and fruit of tree species, prepared by Mary Robeson Sargent (Sargent's wife) at Jesup's request. These, in particular, met with high praise: \"The artist has been true to nature, without loss of refined and purely artistic method, a combination almost unknown in what is called a scientific treatment of natural objects. The result is delightful \u2026 many persons will appreciate for the first time the beauty and grace possessed by the flowers and fruits of many of our common forest trees.\"45 For the benefit of individuals wishing to study the woods from a botanical perspective, a corresponding herbarium had been prepared by Charles Faxon, the assistant director and herbarium curator at the Arnold Arboretum, and shipped to the museum that spring. The Jesup Collection was soon described in the press as \"a credit to the city, and a lasting testimonial to the wisdom and public spirit of Facing page: The press lauded the opening of the Jesup Collection in 1885. This engraving by C. Graham appeared in Harper's Weekly shortly after the exhibition opening. COURTESY OF THE AUTHOR 32 Arnoldia 78\/5-6 \u2022 October 2021 the gentleman who caused it to be created.\"46 It was a first step toward Jesup's original ideal, still awaiting not only more species but examples of economic products and additional illustrations to fully represent the American forests. As far as Sargent's objectives, there was also more to come, but scientific visitors had already found it as informative as it was popular. Worthily Housed In its first incarnation, the woods exhibit occupied the lower floor of the Museum, \"in the space between the rows of side cases,\" leading to the observation on opening day that the space \"is too contracted for this use, and the floor has a cluttered appearance which those who recall its original spaciousness and light will regret. Plainly the time has come when a new wing for the Museum is demanded, so that this collection, unique in its scientific and industrial importance, shall have the sweep of an entire floor.\"47 At the time, the logs shared the hall with the collection of mammals, whose curator was critical of the disruption to those displays.48 Sargent, naturally, weighed in, complaining that \"nothing can be worse than the present mixture of mammals & woods.\"49 While there were already long-term plans for additions to the museum's building, Sargent proposed an alternative idea to Jesup: the museum should construct a separate one-story building for the purpose of housing the wood collection and associated forestry resources, including a library and herbarium, and call it the Jesup Building. He wrote to Jesup, \"The whole thing could be put up in a couple of A large cross-section of a Douglas fir (Pseudotsuga menziesii) appears among cases in the American Museum of Natural History's Forestry Hall, shown in 1903. AMNH RESEARCH LIBRARY DIGITAL SPECIAL COLLECTIONS, 42124 Jesup Collection 33 months and you could have your collection in safe quarters where it could never be interfered with by any one & arranged in such a manner that there never could be any danger of its becoming merged or mixed with the other collections.\"50 It is clear that Sargent wanted to resolve some of the fundamental curatorial problems that the collection was already experiencing, but it is also tempting to suppose that Sargent wanted his own museum of woods (and that Jesup would build it for him). That notion was never pursued, but the Jesup Collection did prevail in occupying the lower hall all to itself. A new display was opened to the public on November 15, 1890, revealing 425 species and almost 250 watercolors, arranged in family groups in the cases along each side of the hall.51 While this was seen as an improvement, and many visitors believed the collection actually was complete, Sargent advised Jesup not a year later, \"I don't think that we ought to consider the arrangement as final or that the collection is worthily housed or properly arranged until some radical change is made by which sufficient room for its display can be had.\"52 In 1893, planning began for the construction of the museum's southeast wing, part of the Seventy-Seventh Street facade, the ground floor of which would be dedicated to the wood collection when it was completed in 1895.53 As the new wing took shape and its opening drew closer, there ensued a paramount disagreement (most emphatic and least charitable on the part of Sargent) over plans for the new hall. In a two-page, typewritten response to Jesup's early scheme for cases and general arrangement, Sargent replied vehemently, and disproportionately: \"A good deal of additional work in connection with the Collection has been laid out for me but I confess I do not feel much like undertaking it if the results are to be as bad as you seem to be determined to make them.\" He asserted that his reputation among scientists could suffer if Jesup's plans were followed, concluding, \"This, from my point of view, is the unfortunate thing in the whole matter and why I believe that I have not been treated properly by you.\"54 Jesup wrote out a six-page reply (that he did not send) in which he recounted their previous discussions about the design. He concluded, \"It would be more agreeable to me in meeting with objections from yourself to have them presented to me in a spirit of help and friendliness \u2026 During the many years of our friendship I have exerted myself to please you, and shall continue to do so in any way I can, but I expect consideration at your hands also.\"55 In place of this letter, Jesup sent museum secretary John H. Winser to consult with Sargent in person about the central points of dispute, namely the design of the new cases and the placement of the immense cross-sections of coast redwood (Sequoia sempervirens) and giant sequoia (Sequoiadendron giganteum). In short, Jesup had wanted to include two or more round cases to break up the \"monotony\" of the exhibit, but doing so would have interrupted the botanical order to a degree that Sargent could not tolerate. At the same time, Jesup had arranged to place the cross-sections of the big trees just outside the main hall, on either side of the entrance, in part because of the architectural requirements for supporting them; Sargent was adamant that they should be placed in the center of the hall with the other logs, despite that this arrangement would require structural reinforcement of the floor. Jesup's proposal took into account the flow of visitors, the overall aesthetic, costs, and the physical constraints of the building; Sargent worried most about what other scientists would think of the exhibit and felt that those concerns had not been adequately considered.56 Citing engineering and safety factors, an Executive Committee of the museum resolved the practical question, temporarily, in favor of the original layout.57 Early in 1896, when the specimens were moved into the new hall and the watercolors were hung, the debate subsided, and Sargent's attention turned back to his usual curatorial concerns. Jesup assured the museum's trustees that the lower hall of the new East Wing had been designated for the \"permanent lodgment\" of the wood collection and concluded, \"It is thought that no better plan can be conceived whereby the effectiveness of the exhibit can be increased.\"58 Not surprisingly, however, even this latest arrangement would be revised again as specimens were added to the exhibit, at Sargent's urging, through the early 1900s.59 Cyrus Pringle \u2014 Pacirsc Northwest, Arizona, California, Texas, and Northern Mexico \u2014 \"He made for the Jesup Collection of North American Woods \u2026 a large collection of timber specimens from some of the most inaccessible and digscult regions \u2026 Becoming interested during this journey in the usora of Mexico, he has for the last twelve years devoted himself exclusively to its exploration. During his annual journeys, which have extended over many of the states, he has made large and unrivaled collections \u2026 and has discovered many undescribed genera and species.\" Samuel Buckley \u2014 Southern Texas \u2014 \"Buckleya, a remarkable Santalaceous genus, of which he discovered the usowers and fruit, and which is represented in the usora of America by a graceful shrub of the mountains of North Carolina \u2026 rstly commemorates Buckley's zealous and too little appreciated labors in the cause of science.\" Jesup Collectors \u2014 More than rsfty collectors helped with acquiring, packaging, and sending large wood specimens for the Jesup Collection of North American Woods. fse specimens originated from thirty-two states, along with four Mexican states and one Canadian province. Several collectors were especially prolrsc. fseir general collecting locales are shown on this map of coniferous and deciduous forests, prairies, and treeless regions, created for the 1880 Census of the United States. Charles Sargent often commemorated the careers of collectors in his Silva of North America. fsese excerpts suggest the nature of the collectors' accomplishments. State or province represented in the collection. Charles Mohr \u2014 Gulf Coast \u2014 \"He made his home at Mobile, Alabama. Here for many years he has been a successful manufacturing druggist, and has devoted his spare time to the study of the usora and the natural resources of the state.\" Allen Curtiss \u2014 Florida and Interior Southeast \u2014 \"He has found many plants, including a number of tropical trees, not known in the territory of the United States before his time.\" John Sears \u2014 Northern New York and Eastern Massachusetts George Letterman \u2014 Missouri, Arkansas, and Northeastern Texas \u2014 \"He rsnally in 1869 settled in Allenton, Missouri, a railroad hamlet about thirty miles west of St. Louis \u2026 fse distribution of the trees of this region before Mr. Letterman's travels was little known, and much useful information concerning them was rsrst gathered by him.\" Henry Ravenel \u2014 South Carolina and Georgia \u2014 \"No other American botanist, perhaps, has minutely studied so many forms of the vegetable kingdom as Ravenel, and none has been more respected or beloved.\" 36 Arnoldia 78\/5-6 \u2022 October 2021 Practically Complete As Sargent's early work on the forest census had concluded in 1884, his focus shifted to taxonomically oriented investigations in support of his Silva of North America and other publications. For nearly two decades, the development of the Jesup Collection was synergistic with that work. Sargent never rested in his ambition to add species to the wood collection, even when his practice conflicted with Jesup's financial concerns and with the museum's pragmatic considerations for their curation. As early as April 1883, after more than two full years of collecting effort, Sargent had indicated that there were twenty-one species needed to complete the collection.60 Still, in February 1886, he reported that there were another \"18 or really 19,\" of which several had already been sent for.61 Just a year later, he wrote, \"I find that there are still a few species which must be added to the Jesup Collection in order to make it complete, and that, moreover, a few important species are not yet properly represented in the Collection.\"62 Sargent reflected in 1889, \"I consider that the collection is practically complete,\" 63 but that notion was short-lived. Sargent soon organized a special expedition to the West Coast and Arizona in 1891 for several unrepresented species. In January 1894, Jesup reported that Sargent had sent him \"the gratifying assurance\" that the collection \"is now complete\"64\u2014even as Sargent was preparing to leave on another collecting trip to Arizona to support his work on the Silva, resulting in at least one new specimen for the museum.65 In April 1898, another twenty-eight species were called for.66 In May 1900, Sargent wrote to museum secretary J. H. Winser, \"We have been finding a lot more trees in the United States during the last year. None of them are very large but all have a scientific interest.\u2026 Now what I want to know is whether I shall go ahead and use my discretion in obtaining such material as may be necessary to complete the Collection.\"67 A year later, Sargent ordered several more specimens from Arkansas, Texas, and Missouri, and noted, \"I understand there is still a good deal more work to do on the collection before it can be considered complete.\"68 Very late in this process, Sargent occasionally accompanied his requests with a lament, such as, \"If it is not continued, I shall be saved a lot of disagreeable bother and letter-writing.\"69 Jesup at times wondered at the necessity of so many very similar species, the number of duplicate specimens that had been sent, and the many that needed to be replaced over time because of damage or decay. He was also not na\u00efve to the fact that he was often financing Sargent's research by supporting new collecting trips for certain trees, and he once expressed frustration about this habit.70 In a note to himself on the back of one letter, Jesup wrote, \"I wonder when the getting of specimens is going to stop.\"71 Both men were clearly tiring of the work of supervising and organizing the collection, wanting it to be both comprehensive and finished, but Jesup's support continued. Still additional specimens were received at the museum late in 1901,72 but by July 1902, Sargent was again discussing sending a collector for more.73 In 1908, the year of Jesup's death, thirty-five specimens (possibly the last) were added to the exhibit.74 Intelligence, Technical Knowledge and Enthusiasm While Sargent continued to direct the collection of new specimens, the opening of the museum's public exhibit in 1885 had added an informal duty: the role of absentee curator. Although S. D. Dill, an experienced carpenter, had been hired specifically to oversee the preparation and installation of the logs and related materials, as well as to build the cases for them, Sargent had ideas of his own about how the collection should be handled and displayed. Beyond persistently lobbying for more space, he involved himself in the minutiae of how logs should be arranged, directly supervised the preparation of labels, and critiqued the display of illustrations following his occasional visits to New York. Only months into the exhibition, Sargent wrote to Jesup with concerns that some specimens housed in new cases were \"already suffering from extremes of temperature as I feared that they would.\" He added that he was \"very anxious & troubled\" that Dill's workroom in the Arsenal was inadequately heated and exposed the specimens to \"danger of destruction by fire or at the hands of outsiders.\"75 Nearly fifteen years later, he offered a similar assessment and insisted that Dill be provided with a workspace Jesup Collection 37 Cross-sections of giant sequoia (Sequoiadendron giganteum, left) and coast redwood (Sequoia sempervirens) flank the entrance to Forestry Hall. The giant sequoia is the only specimen from the Jesup Collection now displayed at the museum. AMNH RESEARCH LIBRARY DIGITAL SPECIAL COLLECTIONS, 5299 that better protected the specimens, adding, \"The money value and cost of these specimens is small in proportion to the expenditure of intelligence, technical knowledge and enthusiasm necessary to procure them, and it is discouraging after all the labor which has been expended in getting them if they are allowed to go to ruin in the Museum.\"76 Although work remained to be done, and to Sargent's dismay, Dill, the collection's chief preparator, caretaker, and de facto on-site curator for twenty years, left the museum for his native Nova Scotia in 1902. To facilitate interpretation of the specimens, museum director Herman C. Bumpus began an inventory of the wood collection in 1903 77 and enlisted Roy W. Miner from the Department of Invertebrate Zoology for the task. Even at that time, the museum's growing bias toward other facets of natural history, to the neglect of botany, was apparent to Bumpus, who frankly acknowledged the economic entomology and wood collections as the entirety of the museum's botanical holdings.78 The \"Forestry Department\" (comprising essentially the collection itself) was without a dedicated curator until 1907, when Alfred C. Burrill, an entomologist by training, was appointed to oversee the exhibit of woods.79 In 1909, Mary C. Dickerson was hired as curator of the Department of Woods and Forestry and served in that capacity for a decade.80 During her editorship of the American Museum Journal, forestry was several times a featured topic. In her 1910 guide to \"Trees and Forestry,\" 38 Arnoldia 78\/5-6 \u2022 October 2021 which drew examples from the Jesup Collection, she expanded on themes of ecology and conservation that were not only current but had long been advocated by the collection's progenitors, Jesup and Sargent.81 Just two years after Jesup's death, museum president Henry F. Osborn reported, \"The Jesup Collection of North American Woods is being rearranged and installed in a way to bring out more clearly the classification of trees, their relationship and their economic uses.\"82 With the wood collection numbering 505 specimens on display, additions were made for several more years in the form of watercolors, photographs, and wax models of foliage, flowers, and fruit;83 Mary Sargent had continued to add to the watercolor series, until more than four hundred paintings were on display with the logs. Space continued to be a problem as time went on (there, and throughout the museum), and activity centered around rearranging specimens to avoid crowding to the extent that was possible.84 Aside from Sargent, who had contributed his knowledge during the collection's genesis, only an oversight committee\u2014chaired in absentia by Gifford Pinchot (cofounder of the Yale Forest School) and James W. Toumey (the school's first Morris K. Jesup Professor of Silviculture)\u2014 afforded forestry expertise after the turn of the century. It was not until 1917 that the department had the benefit of an in-house, credentialed forester. During an era of very limited departmental budget, Yale graduate and future forest ecologist Barrington Moore had been hired as assistant curator, and it was hoped that his experience would contribute to topical research and education at the institution.85 He was shortly called to service in the First World War, however, and by 1920 both he (for other opportunities) and Dickerson (for health reasons) had left the museum. This loss of expertise and energy only compounded the obstacles faced by the wood collection and related subjects that Jesup had promoted. As institutional memory of the collection's formation had been episodically lost since the turn of the century, and the collection's place of priority eroded after the death of its creator and benefactor, its fate became inexorably linked to that of the department going forward. An Old-Fashioned Systematic Arrangement Unlike other collections and exhibits prepared by the various dynamic and actively growing departments of the museum\u2014especially Mammalogy and Ornithology, Paleontology, and Anthropology\u2014the wood collection remained little changed from the 1910s through the 1930s. While the curatorship went unfilled, the Jesup Collection had a champion in museum director Frederic A. Lucas, who in 1922 wrote to President Osborn, \"It is extremely important that we should revive our forestry department, for its own sake and also in memory of Mr. Jesup.\"86 Following Lucas's death in 1929, George H. Sherwood, as museum director and curator of the Department of Education, became its defender. After his death eight years later, the scientific staff of the museum proposed that \"an attempt be made to place some one in charge of the wood collection.\"87 For another decade, the Department of Forestry and Conservation was again chaired and staffed by scientists borrowed from other departments, until a curator was hired for the position in 1946. In the meantime, the finished logs not only occupied an entire exhibit hall but myriad smaller duplicates and miscellaneous wood samples took up valuable storage space when lack of such space at the museum was a chronic problem. Discussions about disposing of the Jesup Collection began to stir at least as early as 1937, when museum director Roy C. Andrews (Sherwood's successor) had suggested that the collection be donated to the New York Botanical Garden \"or some other institution\" in order to create space for new exhibitions. In response, the museum's Council of the Scientific Staff resolved that the collection remained important scientifically as well as to the work of the Department of Education, and argued that to give away this \"superb gift\" could discourage other donations to the museum.88 When the question resurfaced in 1942 under the museum's new director, Albert E. Parr, calls to abandon the wood collection were again met with protest. Informal opinions attributed to the museum's Advisory Committee on Plan and Scope included regret \"that serious proposals have been made to burn up the collecJesup Collection 39 tion,\" and indicated a strong consensus that the museum had an obligation to find \"a satisfactory or a better home for it\" in order to avoid a \"gross\" breach of trust.89 Parr's plans for the museum were dampened during the ensuing years of the Second World War as the institution adjusted to extended absences among curatorial and administrative staff who had joined the armed forces, changes in visitation and patronage, curtailed research activity, and altered demands on the museum's technical and human resources.90 Following the war, Parr discussed the process of \"reconversion\" from the distorted wartime operations of the museum to a post-war vision for its future. He made it clear that he saw this process, both inevitable and necessary, as an opportunity to focus the museum's scope and actively integrate its research and educational activities across disciplines and into the wider landscape of public consciousness. He wanted to find alternatives to standard approaches to exhibition, where \"an old-fashioned systematic arrangement of specimens, unrelieved by an occasionally freer use of artistry, becomes dull and boring to the spectator.\"91 Abandoning staid practices was the foundation for planning the museum's \"program of modernization\" in the years to follow.92 In addition to its orphan status among the departments of the museum, there may have been no single display in the museum at that After more than sixty years on public display, the Jesup Collection was dismantled in Forestry Hall in 1948. AMNH RESEARCH LIBRARY DIGITAL SPECIAL COLLECTIONS, 2A1316 40 Arnoldia 78\/5-6 \u2022 October 2021 time that so epitomized a nineteenth-centurystyle exhibit than the Jesup Collection of North American Woods. Shortly after Parr became the museum's director in 1942, he initiated discussions with botanist Bror E. Dahlgren, once an assistant curator in the Department of Invertebrate Zoology at the museum, who since the 1920s had been affiliated with the Field Museum of Natural History in Chicago. Dahlgren was asked to reconsider how the subjects of forestry and conservation would be represented at the museum. Initially, his advice pertained to a rearrangement of the existing log specimens, \"to break up the single linear, traditional systematic arrangement,\" emphasizing instead the geographic distributions and associations of the many species represented. He envisioned this new scheme as representing the composition and structure of regional American forests, resulting in displays that were more like the dioramas familiar from the museum's zoological exhibits.93 Even with this new thinking toward repurposing the logs, however, the collection's future was not secure. In July 1946, botanist Henry K. Svenson became chair and curator of the reconstituted Department of Forestry and General Botany, which counted two other museum associates, Clarence Hay (anthropology) and Charles Russell (education), as its scientific staff. As a longtime consultant to the museum while a curator at the Brooklyn Botanic Garden, Svenson had AMNH RESEARCH LIBRARY DIGITAL SPECIAL COLLECTIONS, 322587 Artists create detailed replicas of trees for the Olympic Forest diorama during the renovation of the Forestry Hall in 1952. Jesup Collection 41 been designing a new forestry hall and began his tenure at the museum with a preliminary plan for the new exhibits. He recognized the historical importance of the wood collection as \"a heritage of the America that is past, and that our forests would no longer provide such a fine assemblage of material,\" and noted that it would \"become of greater and greater value as time goes on.\" At the same time, Svenson recognized that the future of the department would be a departure from its past. The emphasis of its work would not be on specimens, which would be kept \"behind the scenes,\" but on illustrating the integrated relationships and landscape processes represented by forest vegetation.94 Toward this end, the existing Hall of Forestry was closed on November 1, 1948, after which the exhibits were dismantled.95 As exhibits were revised, Parr explained in 1951 that the role of natural history museums in the progress of science had been evolving over the prior decade. There remained an abiding interest in individual organisms, which were the realm of basic research and a staple of the museum's scientific program. At the same time and increasingly, the museum identified new objectives for their work: understanding the interactions of organisms with their environment (their ecology) and recognizing the necessity for their conservation in nature. It was in these areas where Parr saw the museum's most critical educational mission.96 An early expression of this philosophy was the Felix M. Warburg Memorial Hall of Ecology. Occupying the space where the Jesup Collection had been exhibited, several new exhibits were intended to illustrate the ecosystems of New York State and how the human population influenced the landscape. Adjacent to this, in the southeast corner of the first floor (formerly known as Darwin Hall or the Hall of Invertebrate Zoology), the new Hall of North American Forests was unveiled on May 14, 1958, featuring life-sized dioramas of eleven forest types from across the continent. Where the hundreds of individual trunk segments, separate models of foliage and flowers, and illustrations that populated the former hall had left their forests of origin to the imagination of visitors, the new displays revealed integrated forest ecosystems, with characteristic herbaceous plants, animals, and physical elements (sunlight, water, soils) conspicuously represented in three dimensions. The focus of the new hall was on forests as habitats, the interrelationships among organisms that live in forested regions, and the importance of maintaining these ecosystems.97 Although the tree species themselves were no longer the raisons d'\u00eatre of the new exhibits, the new hall was, effectively, an embodiment of the ideals that its namesake had hoped to promote through the assembly of the original Jesup Collection. The new exhibits were met with admiration.98 Of all the pieces formerly on display, only the large cross-section of giant sequoia remained, as it does today. Meanwhile, as the penultimate step toward disposition, the woods had been officially designated a \"scientific storage collection\" in 1953, and the specimens were sequestered elsewhere in the museum.99 Ponderous and Not Easily Handled In September 1956, Parr ultimately succeeded in convincing the museum's Management Board that \"there was no probability of this material [the wood collection] ever being put to any real use by The American Museum of Natural History.\" He asked the board to approve the transfer of the Jesup Collection to the Smithsonian Institution, which he hoped \"would guarantee proper care and use of the material in accordance with the purposes for which it was collected.\"100 With the board's approval to pursue disposition, then-curator of the museum's Department of Vegetation Studies, Jack McCormick, initiated correspondence with the National Museum to effect this transfer. Because the Smithsonian was preoccupied with the construction of new buildings and other exhibits, these discussions proceeded intermittently over the next two years. The director of the Smithsonian's Museum of Natural History, Remington Kellogg, finally submitted a formal request to Parr in December 1957. His proposal outlined a dramatic new vision for the specimens: Our plans foresee the utilization of the collection in several ways. The large redwood, Sitka spruce, Douglas-fir, sugar pine, ponderosa pine, white pine, oak, walnut, and longleaf pine trunk 42 Arnoldia 78\/5-6 \u2022 October 2021 specimens are being considered in connection with exhibits, in the coming Museum of History and Technology, on early lumbering in the Northeast, the Lake States, the Central Hardwood Region, the Southern Pinery, the Pacific Northwest, and the California Redwood Region. A few of the other large specimens may possibly be halved lengthwise, one half being exhibited with tangentially and radially cut boards from the other half, and the remainder cut into study samples for distribution to educational institutions, colleges, universities, and museums. The remainder of the collection would eventually be cut into study samples for distribution as stated above. We would retain at least two specimens of each species that is cut.101 Parr expressed reticence toward the Smithsonian's plans to destroy the majority of the logs, but he was steadfast in his determination to relocate the huge collection.102 The museum's Board of Trustees approved the transfer at its April 1958 meeting.103 Despite this progress, the arrangements for the collection's transfer remained suspended for another two years. Parr retired, and James A. Oliver became the museum's new director in 1959. During this same time frame, both the directorship of the Smithsonian's Museum of Natural History and the curatorship of its Department of Botany (which included its wood collection) also changed. In 1960, William L. Stern became the Smithsonian's new curator of the Division of Woods. Stern, formerly the curator of the Samuel J. Record wood collection at Yale University, had earlier in that role declined the museum's offer of the Jesup Collection. He explained to McCormick, \"We refused on the grounds that the space needed for storage would be beyond our means, that many of the pieces were ponderous and not easily handled.\" At the Smithsonian, Stern was again faced with the prospect of acquiring the Jesup Collection. In January 1960, he noted to McCormick, \"If I had been Curator of the Division of Woods in the National Museum at the time the Jesup Collection was offered, I do not know how I would have reacted to the offer.\u2026 I just hope that there will be no restrictions on cutting the specimens and that there are no qualifications regarding the handling of the material once it is in the National Museum.\"104 Stern had expressed his opinion to the Smithsonian's new director of the Museum of Natural History, Albert C. Smith, that despite \"the historical importance and unique nature\" of the Jesup Collection, \"it would not greatly increase the usefulness of our present collections for anatomical study.\"105 In his correspondence with Oliver in June 1960, Smith explained, \"One of the problems that we both inherited, in connection with our new positions, concerns the Jesup Collection of Woods of the United States.\u2026 I am now in the embarrassing position of having to ask you to allow the Smithsonian Institution to reverse itself, as to acceptance of the Jesup Collection.\" 106 He indicated that although one or two of the monumental cross-sections might still be useful in their exhibits, the costs of relocation and the ever-present problem of storage were obstacles to their previously agreed-upon plans. Oliver, of course, was disappointed but acknowledged the Smithsonian's position.107 For the sake of the logs, it was certainly a fortuitous development: the very scope and volume of the collection that had inspired museum visitors had made it difficult to accommodate elsewhere, and just as onerous to cut up into tiny hand samples. These were only the first obstacles the museum encountered in its efforts to dispose of the Jesup Collection, but the reasons would not change going forward. McCormick next approached William C. Steere, director of the New York Botanical Garden. After initially suggesting that the garden could accept the Jesup Collection, however, the offer was declined later in 1961.108 Following McCormick's departure from the museum in August of that year, at which time the Department of Vegetation Studies disappeared forever, Oliver took up the cause himself. To an inquiry from Stanley A. Cain, of the University of Michigan School of Natural Resources, he wrote: \"This collection is really a very important one and it should be transferred to a single institution intact. The bulk of the collection is one of the big problems that hinders anyone from accepting it. However, there are no restrictions on it and the wood samples could easily be cut up for other institutions.\"109 This Jesup Collection 43 latest offer was not pursued. With essentially the same preamble, Oliver next approached the Field Museum of Natural History, but received no favorable reply.110 Happy to Turn it Over As Oliver's frustrated efforts began to resemble desperation, a promising inquiry arrived from the Pacific Northwest. Early in 1963, Oliver had spoken with a man named Lloyd S. Millegan, a retired public servant who lived in McMinnville, Oregon, and ran a small marquetry business, Lloyd's of Oregon, in nearby Portland. Millegan envisioned mounting a display of the logs at the New York World's Fair in 1964, then displaying the collection in Portland to generate publicity and business for his handicrafts. Having been unsuccessful in finding another museum to accept the collection, Oliver explained that the museum was \"eager\" and \"would be happy to turn it over to anyone who will undertake the cost of packing and transporting the entire collection from the museum to the new location.\" He emphasized that \"the entire collection be taken in its entirety because we have no personnel to dispose of it properly piecemeal.\" 111 When another group, coincidentally also in Portland, inquired about the collection later that year, Oliver asked Millegan to submit a formal offer indicating his intentions and to confirm that the collection would be removed by February 1964.112 While Oliver awaited word from Millegan, he continued to entertain correspondence with Aldred A. Heckman, director of the Louis W. and Maud Hill Family Foundation in St. Paul, Minnesota. Through the common acquaintance of William Steere at the New York Botanical Garden, the Hill Family Foundation had been in discussions with the Gallery of Trees Committee, a group of industry and civic leaders as well as forestry professionals, about assisting them in acquiring the Jesup Collection for their museum in Portland. Heckman explained, \"There is real interest in having the Collection in Portland.\" He emphasized that there was both local expertise available to prepare and interpret the proposed exhibit, as well as an audience already interested in trees and forestry attending the existing forestry museum. Further, the City of Portland and the Oregon Museum of Science and Industry had indicated willingness to participate in structuring the acquisition.113 Steere himself wrote to Heckman, \"Naturally I am deeply grateful to you for your personal interest in seeing that an exhibit of national importance is not reduced to veneer or small samples\u2014or ashes.\"114 At an early meeting in January 1964, the Gallery of Trees Committee proceeded to address questions about transportation of the collection and the siting, design, and construction of a new building to house it. The Hill Family Foundation offered to defray the costs of transporting the collection to Portland, provided that it be publicly owned and exhibited. The City of Portland's Park Bureau and the Oregon Museum of Science and Industry were identified as the preferred partners.115 Whether it had intended to or not, the meeting illustrated the contrast between the committee's plans, for which the organizers could demonstrate institutional, technical, intellectual, and financial support, and those of Millegan, whose intentions had not addressed any of the real practicalities involved with adopting these specimens. Both the Gallery of Trees Committee and the Hill Family Foundation had been surprised to learn of Millegan's prior claim, but their strong interest in obtaining the logs for Portland's museum compelled them to include him in their discussions. Millegan was asked to explain his relationship to the collection. The meeting minutes recorded: \"He asked for it not knowing then what could be done with it. His offer was accepted.\u2026 [He] said he had no deed for the collection, merely a letter saying he could have it.\"116 He was asked what conditions he would place on forfeiting his \"claim\" to the collection so that the committee could proceed. Millegan stipulated first that the collection should be freely accessible and well presented; beyond that, he wanted to use the exhibit to educate visitors about marquetry and its use of various woods, and to display his marquetry products alongside the exhibit.117 At this time, Heckman indicated to Oliver that there would be no further discussion among the foundation and the entities in Portland until Millegan's position was clarified. He concluded, \"It seemed 44 Arnoldia 78\/5-6 \u2022 October 2021 to me that we were rapidly getting to the point of having too many cooks as far as the North American Woods Collection is concerned.\"118 The chair of the Gallery of Trees Committee, Thornton T. Munger, addressed Oliver shortly after the meeting, indicating that the committee was \"impatient\" to understand where they stood in relation to Millegan's plans to acquire the collection.119 Heckman soon wrote to Oliver, as well, reinforcing the message of progress that had been made toward planning for the collection's move to Portland under the assumption that Millegan would cede the collection. He added, \"We thought that if funds were assured to cover the costs of transporting the Collection to Portland and preparing it for display, the decisions regarding these other matters would be made with reasonable speed. This is as far as we can go. The next steps will have to be taken in Portland.\"120 Millegan subsequently contacted the committee to revise his terms for relinquishing his claim to the collection, introducing the demand that he be allowed \"to operate in the exhibit area a concession where selected gift and educational items in wood could be purchased.\" The committee's chair, Munger, was a retired forester of long tenure in the U.S. Forest Service whose career and research had been devoted to developing methods for sustainable forestry and conservation. He and the Gallery of Trees Committee envisioned a much broader mission for the collection, that it would illustrate the forest resources of the country for the benefit of public education. Neither the committee, nor the City of Portland, nor the Hill Family Foundation approved of the idea of using the collection to support a commercial enterprise, which in terms of the proposed new building would also be prohibited by city ordinance.121 Although the committee was at an impasse as the negotiations stretched into April, May, and June, Munger had continued to plan as though a compromise would eventually be reached.122 After hearing again from Munger following a meeting in May, Oliver decided to finally draw the matter to a close. He informed Millegan in June, \"You have repeatedly stated that you were interested in acquiring this collection and were given several deadlines for the acquisition of the collection.\u2026 I think we have been exceedingly patient in waiting for you to fulfill your intentions. Therefore, your option to the collection has been withdrawn and we shall seek to dispose of the collection through other channels.\" 123 Oliver notified Munger of the transaction and renewed his offer to the Gallery of Trees Committee, with the only requirement being \"that we hope it will be exhibited for the benefit of the public and will be available to students for study.\" He urged that the collection be transferred by September 1.124 The Gallery of Trees Committee was relieved, the Hill Family Foundation was satisfied, and the City Council and Oregon Museum of Science and Industry all agreed that the collection would finally belong to Portland. In the meantime, the Gallery of Trees Committee had reached a consensus about the location for the new exhibit. Rather than constructing a new building, the Jesup Collection could be displayed on the unoccupied second story of the old Forestry Building, a stupendous log structure that had been built in northwest Portland for the Lewis and Clark Centennial Exposition of 1905. The main floor was already in use as a museum of forestry and the logging industry, and it was thought that the log specimens would complement these exhibits. Because the aging balconies required engineering changes to accommodate the collection, the committee intended to store the collection once it arrived in Portland while funding was raised for the renovations.125 Just a month after the final July meeting that approved of these plans, tragedy swept them all aside. A fire started in the office of the Forestry Building on the evening of August 17 and rapidly spread to the entire structure. The next morning, Munger observed the smoldering remains, which included the entire contents of the city's forestry museum that he had helped to oversee.126 By 1971, when the new Western Forestry Center building opened, the story of the calamity in the museum's own informational materials had come to include the Jesup Collection and its miraculous escape of this fate by having still been in storage in Portland.127 Twenty years after the fire, the story read: \"When the old log museum burned in August Jesup Collection 45 1964, two box cars full of the Jesup collection had just arrived. Sidetracked and waiting to be unloaded, the collection narrowly missed destruction in the fire. The exhibit then was stored by the city until the new forestry center opened in June 1971.\"128 In fact, the Jesup Collection had still been safely in New York. Munger wrote to Oliver just days after the fire, expressing the committee's sadness at the loss and explaining its plans to rebuild. He noted, \"It is very fortunate that the Jesup Collection was not there.\"129 At the museum, Oliver and his staff were solidifying plans for an early October moving day. The Santini Brothers moving company was contracted to pack and transport the collection.130 On October 6, 1964, the specimens departed the museum aboard three moving vans destined for Portland, Oregon (the surviving paperwork gives no indication that railroad cars were employed).131 How they were stored once they arrived there is not recorded, but it is possible that the Gallery of Trees Committee took advantage of one of the offers for local warehouse space that had been made during their planning process.132 The Jesup Collection would not be put on display for nearly seven more years while a new building was constructed, but that building promised to include dedicated space for the logs. At the new Western Forestry Center, which opened in June 1971 in Washington Park, west of downtown Portland, the Jesup Collection In 1971, the Jesup Collection of Woods reopened in a new home at the Western Forestry Center in Portland, Oregon. ARCHIVES OF THE ARNOLD ARBORETUM 46 Arnoldia 78\/5-6 \u2022 October 2021 was \"the background theme that links together feature displays at the Forestry Center. Some of the largest logs are stationed at the entrance and around the outdoor covered walkway; inside, smaller specimens circle the first-floor display room. Other logs fill corners and file along corridors.\" 133 Following their move, the logs had been cleaned, refinished, and given new labels by local members of the Society of American Foresters and the International Wood Collectors Society. The historical value of the 505 logs said to be on display, representing trees of such stature that in many cases could no longer be observed in the United States, was well appreciated, and the collection remained a popular exhibit.134 As the Western Forestry Center expanded its educational mission and shifted its focus to forests at a global scale, taking on the name World Forestry Center in 1986, the collection's relevance was again eclipsed by its physical footprint. About January 1994, the collection was donated to Agricenter International in Memphis, Tennessee.135 Although exhibited there for several years, the logs have since spent more than two decades in storage. A Heritage Following Jesup's death, Sargent reflected, \"The formation of the Jesup collection of North American Woods \u2026 was a matter of national importance. The preparation of this collection enabled us to study the distribution of the economic value of many trees which, before Mr. Jesup's undertaking, were largely unknown. I think it can be said that this collection is the finest representation of forest wealth that exists in any country.\"136 In its time on exhibit, the collection was marveled at by audiences for more than eighty years altogether. It provided not only Jesup and Sargent but some early influencers of American forestry\u2014including Heinrich Mayr, Carl A. Schenck, Gifford Pinchot, Bernhard E. Fernow, Barrington Moore, and later even Thornton Munger\u2014with inspiration and a platform to promote a growing movement supporting the conservation of American forests. What the logs represent has not changed, and their historical significance has only grown. Apart from the varied circumstances leading to their assembly in New York from all across North America, as a group the collection has twice crossed the country; it has evaded annihilation more than once, each time saved by wellmeaning caretakers facing formidable logistical challenges. More than 120 years since the consolidation of the collection, although many of the logs are superficially weathered and show wear and tear from handling and the elements, their number is mainly intact. The wood itself has largely not suffered and will be restorable in some future, truly permanent, home. Research to document the geographic origins of individual logs is ongoing; these findings will enable many of them to retake their scientific potential, where study of the wood itself may contribute meaningfully to the knowledge of our environmental past. All of them may yet function as emissaries for their species and for the forested regions from which they came\u2014 possibly even more so today than at the time of the collection's unveiling, when many contemporaries believed that such trees would be lost from America's forests in time, even as forests generally were disappearing, and that such a collection could never again be made.137 Acknowledgments This research was supported in part by a 2019 Sargent Award for Visiting Scholars from the Arnold Arboretum of Harvard University. For their assistance, the author is grateful to the curators of the Harvard University Herbaria; Lisa Pearson at the Arnold Arboretum; Rebecca Morgan and Gregory Raml at the Archives of the American Museum of Natural History; Alex Wiedenhoeft and Regis Miller at the USDA Forest Products Laboratory, Madison, Wisconsin; John Butler and John Charles Wilson at Agricenter International, Memphis, Tennessee; and Mark Reed, Beavercreek, Oregon. Notes 1 A Noble Gift, Sun (New York), 17 May 1885, p.8. 2 American Woods, Harper's Weekly 29(30 May 1885), p.350. 3 Brewer, 1877: 4. 4 Norton, 1879: 110. 5 A. S. Bickmore to C. S. Sargent, 12 Sep 1880, Letterpress Books, 3a: 273, AMNH. 6 Joseph H. Choate, in Hovey 1907: 5. 7 C. S. Sargent to M. K. Jesup, 2 Nov 1885, Departmental Records, 091, Ser. I, AMNH. 8 C. S. Sargent to M. K. Jesup, 11 Nov 1880, Departmental Records, 091, Ser. I, AMNH. 9 Trustees of the American Museum of Natural History 1881. 10 C. S. Sargent to M. K. Jesup, 14 Dec 1880; C. S. Sargent to J. J. Bargin, 20 Apr 1881, Departmental Records, 091, Ser. I, AMNH. Jesup Collection 47 11 M. K. Jesup to C. S. Sargent, 12 Mar 1881, Letterpress Books, 4: 19, AMNH. 12 A. S. Bickmore to C. S. Sargent, 29 Jun 1881, Letterpress Books, 4: 55; C. S. Sargent to M. K. Jesup, 5 Jul 1882, Departmental Records, 091, Ser. I, AMNH. 13 M. K. Jesup to C. S. Sargent, 1 Sep 1881, Letterpress Books, 4: 80, AMNH. 14 M. K. Jesup to A. S. Bickmore, 1 Sep 1881, Letterpress Books, 4: 82, AMNH. 15 C. S. Sargent to M. K. Jesup, 25 Apr 1881, Departmental Records, 091, Ser. I, AMNH. 16 C. S. Sargent to M. K. Jesup, 18 Feb 1896, Departmental Records, 091, Ser. I, AMNH. 17 C. S. Sargent to M. K. Jesup, 20 Aug 1881, Departmental Records, 091, Ser. I, AMNH. 18 C. S. Sargent to M. K. Jesup, 25 Apr 1881, Departmental Records, 091, Ser. I, AMNH. 19 M. K. Jesup to C. S. Sargent, 21 Jun 1882, Letterpress Books, 4: 277; J. J. Bargin to M. L. Saley, 3 Mar 1886, Letterpress Books, 9: 83, AMNH. 20 C. S. Sargent to M. K. Jesup, 2 Nov 1885, Departmental Records, 091, Ser. I, AMNH. 21 C. S. Sargent to M. K. Jesup, 25 Apr 1881, Departmental Records, 091, Ser. I; A. S. Bickmore to C. S. Sargent, 17 Aug 1881, Letterpress Books, 4: 73, AMNH. 22 C. S. Sargent to J. H. Winser, 17 Jun 1899, Departmental Records, 091, Ser. I, AMNH. 23 M. K. Jesup to C. S. Sargent, 28 Apr 1881, Letterpress Books, 4: 35, AMNH. 24 Sargent, 1884. 25 A. S. Bickmore to C. S. Sargent, 3 Dec 1881, Letterpress Books, 5: 50, AMNH. 26 J. J. Bargin to C. S. Sargent, 10 Apr 1882, Letterpress Books, 4: 235, AMNH. 27 Osborn, 1911. 28 The Woods of America\u2014A Great Collection of 394 Specimens, New York Times, 22 Oct 1882, p.13. 29 A. S. Bickmore to C. S. Sargent, 17 Aug 1881, Letterpress Books, 4: 72, AMNH. 30 S. D. Dill to M. K. Jesup, 8 Nov 1882; C. S. Sargent to M. K. Jesup, 1 Jun 1884, Departmental Records, 091, Ser. I, AMNH. 31 J. H. Winser to M. K. Jesup, 26 Jan 1899, Departmental Records, 091, Ser. I, AMNH. 32 C. S. Sargent to M. K. Jesup, 9 May 1882, Departmental Records, 091, Ser. I; M. K. Jesup to C. S. Sargent, 16 May 1882, Letterpress Books, 4: 250, AMNH. 33 Osborn, 1911: 19. 34 S. D. Dill to J. J. Bargin, 24 Oct 1882, Departmental Records, 091, Ser. I, AMNH. 35 C. Vaux to Department of Public Parks, 13 Nov 1882, Early Admin Files, CN1739, AMNH. 36 C. S. Sargent to M. K. Jesup, 1 Nov 1882, Departmental Records, 091, Ser. I, AMNH. 37 Trustees of the American Museum of Natural History 1883: 6. 38 American Wood Specimens: Mr. Jesup's Present to the Museum of Natural History, New York Times, 26 Dec 1883, p.8. 39 C. S. Sargent to M. K. Jesup, 15 Feb 1884, Departmental Records, 091, Ser. I, AMNH. 40 C. S. Sargent to M. K. Jesup, 30 Apr 1884; C. S. Sargent to A. S. Bickmore, 7 Jun 1884, Departmental Records, 091, Ser. I, AMNH. 41 Bickmore, 1885: 778-779. 42 Sargent, 1885. 43 C. S. Sargent to M. K. Jesup, 26 Apr 1885, Departmental Records, 091, Ser. I, AMNH. 44 Jesup, 1885: 6-7. 45 A Noble Gift, Sun (New York), 17 May 1885, p.8. 46 American Woods, Harper's Weekly 29(30 May 1885), p.350. 47 The Jesup Collection\u2014All the Woods of the United States, Formal Opening To-day at the Museum of Natural History, New York Times, 18 May 1885, p.1. 48 J. J. Bargin to M. K. Jesup, 9 Oct 1885, Departmental Records, 091, Ser. I, AMNH. 49 C. S. Sargent to M. K. Jesup, 13 Oct 1885, Departmental Records, 091, Ser. I, AMNH. 50 C. S. Sargent to M. K. Jesup, 13 Oct 1885, Departmental Records, 091, Ser. I, AMNH. 51 An Interesting Collection\u2014Mr. Jesup's Gift to the American Museum of Natural History, New York Times, 16 Nov 1890, p.9; Sargent 1890b; Jesup 1891. 52 C. S. Sargent to M. K. Jesup, 9 Jun 1891, Departmental Records, 091, Ser. I, AMNH. 53 Jesup, 1894. 54 C. S. Sargent to M. K. Jesup, 2 Jan 1895, Departmental Records, 091, Ser. I, AMNH. 55 M. K. Jesup to C. S. Sargent, 17 Jan 1895, Departmental Records, 091, Ser. I, AMNH. 56 J. H. Winser to M. K. Jesup, 23 Jan 1895, Departmental Records, 091, Ser. I, AMNH. 57 Extract of Minutes, Regular Meeting of the Exectutive Committee, 20 Dec 1895, Central Archives, 1203, AMNH. 58 Jesup, 1896: 14. 59 Jesup, 1898, 1899, 1907. 60 C. S. Sargent to M. K. Jesup, 6 Apr 1881, Departmental Records, 091, Ser. I, AMNH. 61 C. S. Sargent to M. K. Jesup, 19 Feb 1886, Departmental Records, 091, Ser. I, AMNH. 62 C. S. Sargent to M. K. Jesup, 11 May 1887, Early Admin Files, CN2169, AMNH. 63 C. S. Sargent to B. Strong, 12 Feb 1889, Departmental Records, 091, Ser. I, AMNH. 64 Extract of Minutes, Regular Meeting of the Exectutive Committee, 19 Jan 1894, Central Archives, 1203, AMNH. 65 C. S. Sargent to M. K. Jesup, 13 Sep 1894, Departmental Records, 091, Ser. I, AMNH. 66 C. S. Sargent to J. H. Winser, 21 Apr 1898, Departmental Records, 091, Ser. I, AMNH. 67 C. S. Sargent to J. H. Winser, 29 May 1900, Early Admin Files, CN3540, AMNH. 68 C. S. Sargent to J. H. Winser, 26 Oct 1901, Departmental Records, 091, Ser. I, AMNH. 69 C. S. Sargent to J. H. Winser, 7 Jun 1899, Departmental Records, 091, Ser. I, AMNH. 70 C. S. Sargent to J. H. Winser, 7 May 1898, Departmental Records, 091, Ser. I, AMNH. 71 C. S. Sargent to J. H. Winser, 1 Nov 1898, Departmental Records, 091, Ser. I, AMNH. 72 C. S. Sargent to J. H. Winser, 4 Dec 1901, Departmental Records, 091, Ser. I, AMNH. 73 C. S. Sargent to J. H. Winser, 23 Jul 1902, Departmental Records, 091, Ser. I, AMNH. 74 Osborn, 1909. 48 Arnoldia 78\/5-6 \u2022 October 2021 75 C. S. Sargent to M. K. Jesup, 6 Oct 1885, Departmental Records, 091, Ser. I, AMNH. 76 C. S. Sargent to M. K. Jesup, 7 Jan 1899, Departmental Records, 091, Ser. I, AMNH. 77 H. C. Bumpus to C. S. Sargent, 14 Oct 1903, Departmental Records, 091, Ser. I, AMNH. 78 H. C. Bumpus to H. H. Kopman, 28 Oct 1904, Departmental Records, 091, Ser. I, AMNH. 79 Osborn, 1908. 80 Osborn, 1910. 81 Dickerson, 1910. 82 Osborn, 1910: 41. 83 Osborn, 1911: 118. 84 Dickerson, 1912. 85 Dickerson, 1917. 86 H. F. Osborn to F.A. Lucas, 11 Dec 1922, Central Archives, 777, AMNH. 87 H. E. Anthony, Meeting Minutes, p.88, Council of the Scientific Staff, 4 Oct 1937, Departmental Records, 086, AMNH. 88 H. E. Anthony, Meeting Minutes, pp.88-89, Council of the Scientific Staff, 4 Oct 1937, Departmental Records, 086, AMNH. 89 Papers presented by the Advisory Committee on Plan and Scope, 6 May 1942, Central Archives, 1232, AMNH. 90 Parr, 1943. 91 Parr, 1946: 13. 92 Davison, 1946: 4. 93 B. E. Dahlgren to A. E. Parr, 6 Jul 1943, Central Archives, 1203, AMNH. 94 H. K. Svenson, Report: Department of Forestry and General Botany, 1947, Departmental Records, 091, Ser. IV, AMNH. 95 AMNH Department of Education Division of Publications 1949. 96 Parr, 1943, 1951. 97 Burns, 1958. 98 Museum Opening Hall of Forests, by S. Knox, New York Times, 14 May 1958, p.35. 99 A. E. Parr, Report of the Management Board, Special Meeting of the Board of Trustees, 20 Apr 1953, Central Archives, 1118, AMNH. 100 Extract of Minutes, Management Board Meeting, 27 Sep 1956, Central Archives, 1203, AMNH. 101 R. Kellogg to A. E. Parr, 19 Dec 1957, Central Archives, 1203, AMNH. 102 A. E. Parr to R. Kellogg, 26 Dec 1957, Central Archives, 1203, AMNH. 103 Extract of Minutes, Spring Meeting of the Board of Trustees, 28 Apr 1958, Central Archives, 1117, AMNH. 104 W. L. Stern to J. McCormick, 15 Jan 1960, Central Archives, 1203, AMNH. 105 A. C. Smith to J. A. Oliver, 24 Jun 1960, Departmental Records, 125, Ser. I, AMNH. 106 A. C. Smith to J. A. Oliver, 24 Jun 1960, Departmental Records, 125, Ser. I, AMNH. 107 J. A. Oliver to A. C. Smith, 5 Jul 1960, Departmental Records, 125, Ser. I, AMNH. 108 J. McCormick to J. A. Oliver, 20 Jun 1961, Central Archives, 1203, AMNH. 109 J. A. Oliver to S. A. Cain, 26 Dec 1961, Departmental Records, 125, Ser. I, AMNH. 110 J. A. Oliver to J. Millar, 26 Dec 1961, Departmental Records, 125, Ser. I, AMNH. 111 J. A. Oliver to L. S. Millegan, 19 Feb 1963, Departmental Records, 125, Ser. I, AMNH. 112 J. A. Oliver to L. S. Millegan, 15 Nov 1963, Departmental Records, 125, Ser. I, AMNH. 113 A. A. Heckman to J. A. Oliver, 15 Jan 1964, Departmental Records, 125, Ser. I, AMNH. 114 W. C. Steere to A. A. Heckman, 17 Jan 1964, Departmental Records, 125, Ser. I, AMNH. 115 Minutes of Meeting to Discuss Prospects of Getting for Portland the Jesup Collection of Woods, 15 Jan 1964, Departmental Records, 125, Ser. I, AMNH. 116 Minutes of Meeting to Discuss Prospects of Getting for Portland the Jesup Collection of Woods, 15 Jan 1964, Departmental Records, 125, Ser. I, AMNH. 117 Minutes of Meeting to Discuss Prospects of Getting for Portland the Jesup Collection of Woods, 15 Jan 1964, Departmental Records, 125, Ser. I, AMNH. 118 A. A. Heckman to J. A. Oliver, 15 Jan 1964, Departmental Records, 125, Ser. I, AMNH. 119 T. T. Munger to J. A. Oliver, 17 Feb 1964, Departmental Records, 125, Ser. I, AMNH. 120 A. A. Heckman to J. A. Oliver, 20 Feb 1964, Departmental Records, 125, Ser. I, AMNH. 121 T. T. Munger to J. A. Oliver, 14 Apr 1964, Departmental Records, 125, Ser. I, AMNH. 122 Minutes of Meeting of Committee on the Jesup Collection of Wood, 29 May 1964, Departmental Records, 125, Ser. I, AMNH. 123 J. A. Oliver to L. S. Millegan, 24 Jun 1964, Departmental Records, 125, Ser. I, AMNH. 124 J. A. Oliver to T. T. Munger, 24 Jun 19","distinct_key":"arnoldia-2021-Such a Fine Assemblage: The Jesup Collection of North American Woods"},{"has_event_date":0,"type":"arnoldia","title":"Such a Fine Assemblage: The Jesup Collection of North American Woods","article_sequence":7,"start_page":24,"end_page":49,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25742","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160af27.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Mauz, Kathryn","article_content":"64, Departmental Records, 125, Ser. I, AMNH. 125 Statement to the City Council of Portland regarding transfer of the Jesup Collection to the Oregon Museum of Science and Industry, 14 Jul 1964, Departmental Records, 125, Ser. I, AMNH. 126 Curator Watches as Embers of History Linger On\u2014 Oregon Logging Associates Consider Plans to Restore Forestry Building, Oregonian, 19 Aug 1964, p.1. 127 Press release: Jesup Wood Collection, Western Forestry Center, Portland, Ore., ca. 1971, I G 9.1 WUS: Woods of the United States Exhibit Records, AA. 128 Jesup Wood Collection historic, enduring exhibit, by J. Sansregret, Oregonian, 28 Sep 1984, p.D7. 129 T. T. Munger to J. A. Oliver, 22 Aug 1964, Departmental Records, 125, Ser. I, AMNH. 130 P. H. Grouleff to J. A. Oliver, 29 Sep 1964, Departmental Records, 125, Ser. I, AMNH. 131 Bill of Lading and Freight Bill, United Van Lines, 6 Oct 1964, Central Archives, 1203, AMNH. 132 T. T. Munger to J. A. Oliver, 2 Jul 1964, Departmental Records, 125, Ser. I, AMNH. 133 Sansregret, 1984: 50. 134 Reed, 1987. 135 Meeting minutes, 25 Jan 1994, Agricenter International (Memphis, Tennessee). 136 Brown, 1910: 165-166. 137 The Woods of America\u2014A Great Collection of 394 Specimens, New York Times, 22 Oct 1882, p.13; Sargent 1890a. Jesup Collection 49 Primary Sources Archival resources have been used with permission and are housed at the Archives of the Arnold Arboretum of Harvard University, Boston [AA]; the American Museum of Natural History Research Library Archives, New York [AMNH]; and elsewhere as indicated. Published Sources Cited AMNH Department of Education Division of Publications. 1949. General guide to the exhibition halls of the American Museum of Natural History. Science Guide 118 (5th ed.). American Museum of Natural History. Brewer, W. H. 1877. General report of the judges of Group VI and report on awards. Pages 1-50 in: Walker, F. A. (Ed.), Reports and awards, Group VI. United States Centennial Commission. J. B. Lippincott & Co. Brown, W. A. 1910. Morris Ketchum Jesup, a character sketch. Charles Scribner's Sons. Burns, W. A. (Ed.). 1958. General guide to the American Museum of Natural History. Science Guide 118 (revised ed.). American Museum of Natural History, Man and Nature Publications. Davison, F. T. 1946. Seventy-seventh report of the President. Annual Report of the American Museum of Natural History, 77: 1-6. Dickerson, M. C. 1910. Trees and forestry: An elementary treatment of the subject based on the Jesup Collection of North American Woods in the American Museum of Natural History. Guide Leaflet 32. American Museum of Natural History. Dickerson, M. C. (Ed.). 1912. Museum notes. American Museum Journal, 12: 35-40. Dickerson, M. C. (Ed.). 1917. Museum notes. American Museum Journal, 17: 76-80. Hovey, E. O. (Ed.). 1907. Introduction: Pioneers of American science: An account of the exercises held and the addresses delivered at the American Museum of Natural History, December 29, 1906. Guide Leaflet 25. American Museum Journal, 7(Supplement): 3-7. Jesup, M. K. 1891. Twenty-second annual report. Annual Report of the Trustees of the American Museum of Natural History ... for the Year 1890-91: 7-13. Jesup, M. K. 1894. Twenty-fifth annual report. Annual Report of the President of the American Museum of Natural History ... for the Year 1893: 7-15. Jesup, M. K. 1896. Twenty-seventh annual report. Annual Report of the President, Act of Incorporation, Contract with the Department of Public Parks, Constitution, By-laws and List of Members for the Year 1895: 7-23. Jesup, M. K. 1898. Twenty-ninth annual report. Annual Report of the President of the American Museum of Natural History ... for the Year 1897: 9-27. Jesup, M. K. 1899. Thirtieth annual report. Annual Report of the President of the American Museum of Natural History ... for the Year 1898: 9-26. Jesup, M. K. 1907. Thirty-eighth annual report. Annual Report of the President of the American Museum of Natural History ... for the Year 1906: 11-34. Norton, F. H. 1879. Illustrated historical register of the Centennial Exhibition, Philadelphia, 1876, and the Exposition Universelle, Paris, 1878. The American News Co. Osborn, H. F. 1908. Thirty-ninth annual report. Annual Report of the American Museum of Natural History, 39: 15-48. Osborn, H. F. 1909. Fortieth annual report. Annual Report of the American Museum of Natural History, 40: 15-42. Osborn, H. F. 1910. Forty-first annual report. Annual Report of the American Museum of Natural History, 41: 15-51. Osborn, H. F. (Ed.). 1911. The American Museum of Natural History: Its origin, its history, the growth of its departments to December 31, 1909 (2nd ed.). New York: The Irving Press. Parr, A. E. 1943. The year's work. Annual Report of the American Museum of Natural History, 74: 5-21. Parr, A. E. 1946. In transition. Annual Report of the American Museum of Natural History, 77: 7-21. Parr, A. E. 1951. Purposes and progress report of the Director. Annual Report of the American Museum of Natural History, 82: 7-36. Reed, M. 1987. The Jesup Collection of Woods. Forest World Magazine, 3(1): 7-11. Sansregret, J. 1984. A history in wood. American Forests Magazine, 90(9): 50. Sargent, C. S. 1884. Report on the forests of North America (exclusive of Mexico). Tenth United States Census, vol. 9. Census Office, Department of the Interior. Government Printing Office. Sargent, C. S. 1885. American Museum of Natural History Jesup Collection: The woods of the United States. D. Appleton and Co. (John Wilson and Son). Sargent, C. S. 1890a. Recent publications: The forests of North America, I. Garden and Forest, 3: 193-194. Sargent, C. S. (Ed.). 1890b. The Jesup Collection of the Woods of the United States. Garden and Forest, 3: 570. Trustees of the American Museum of Natural History. 1881. Twelfth annual report. Annual Report of the American Museum of Natural History, 12: 5-12. Trustees of the American Museum of Natural History. 1883. Fourteenth annual report. Annual Report of the American Museum of Natural History, 14: 5-10. Kathryn Mauz writes from Colorado. Previous publications include a 2018 book on Jesup collector Cyrus Pringle: C. G. Pringle: Botanist, Traveler, and the \"Flora of the Pacific Slope\" (1881-1884).lants imported from Europe and farther afield. William Jr. and his son William Robert Prince took up the cause of identifying and describing plant material so that it could be offered to the public\u2014and they were highly invested in acquiring newly introduced species. In 1804, for instance, Meriwether Lewis and William Clark embarked upon the Missouri River to explore the recently acquired Louisiana Purchase. The expedition had been commissioned at Jefferson's request, and when the explorers returned east, they came bearing seeds and other botanical collections. The Princes were among the first nursery operators to grow and distribute plants from the expedition, and the Oregon grape holly (Mahonia aquifolium) became one of their most successful new products. The Princes were also among the first American nurseries to offer ornamental species from East Asia, like the golden rain tree (Koelreuteria paniculata), lacebark elm (Ulmus parvifolia), and Chinese wisteria (Wisteria sinensis). By the mid-1830s, William Jr. had ten nursery outbuildings, of which several were greenhouses that contained tropical and subtropical plants from Africa and Asia. Visitors could pay an admission fee to experience the warmth and humidity of the greenhouse\u2014a rewarding respite to escape the dark, cold New York winter. The nursery catalogue listed ten tropical hibiscuses (Hibiscus) and two gardenias (Gardenia) that bloomed in their greenhouses. Prince grew tropical fruits and flowers specifically for winter viewing. For variety, they also exhibited insectivorous plants such as sundew (Drosera), pitcher plant (Sarracenia), and Venus flytrap (Dionaea). Moreover, in 1833, The New-York Annual Register reported that the gardens and nursery covered up to forty In 1793, William Prince Jr. purchased twenty-four acres alongside the original nursery, naming the new property the Linnaean Botanic Garden and Nursery. In the decades to come, a cohort of nurseries would open in Flushing, including Parsons Nursery and Bloodgood Nursery, both mapped nearby in 1841. SMITH, 1841\/LIBRARY OF CONGRESS, GEOGRAPHY AND MAP DIVISION ies cultivated in America, other than apples. (While the father and son intended to treat apple cultivation with a third volume, the work was never published.) Like A Short Treatise on Horticulture, this book was widely read in America and became influential among aspiring horticulturalists. Moreover, the Princes paid particular attention to the nomenclature of the fruits covered in all of the publications, untangling confusion occurring in the field. This interest in the accurate classification of horticultural plants began with the work of William Sr., and it was among the family's most significant contributions to American horticulture. As a testament to William Jr.'s interest in classification, he displayed in his home a bust of Carl Linnaeus, the Swedish botanist who formalized the modern system of botanical nomenclature. William Jr. received the statue in a presentation by New York governor DeWitt Clinton at a meeting of European and American scientists to honor Linnaeus's birthday in 1823. A simultaneous celebration in Virginia was officiated by Thomas Jefferson, an honorary member of the Linnaean Society of Paris. By the time William Jr. died in 1842, Flushing had become a vibrant center for American horticulture. Bloodgood Nursery had been established there in 1798 and would become known as a specialist in maples. (A common Japanese maple even bears the name of the nursery: Acer palmatum 'Bloodgood'.) G. R. Garretson Nursery, a seed company specializing in flowers and vegetables, was established in 1836 and would grow to cover one hundred acres, supplying wholesale seeds to nurseries across the United States and offering retail via mail order. But the most famous of these newer operations was Parsons Nursery, established in 1838; the Parsons family would later play a central role in introducing plants from East Asia, especially Japan. Meanwhile, William Robert had been assuming increasing responsibility for the Linnaean Botanic Garden and Nurseries. In the 1820s, he expanded the nursery, purchasing three large parcels so that his land holdings may have totaled up to 113 acres. These properties were located adjacent to a house he bought for himself in 1827. The home had a wide center hall, \u222b 20 Arnoldia 78\/5-6 \u2022 October 2021 acres and contained approximately ten thousand species of trees and plants, with particular attention devoted to grapes and mulberry trees. Visitors had free access to the outdoor gardens every day, except for Sundays. At the same time, the commercial operations of the nursery expanded rapidly, as evidenced by William Jr.'s increasingly thicker plant catalogues. He also began to subdivide the products among smaller specialized catalogues. In addition to his standard Annual Catalogue for Fruit and Ornamental Trees and Plants, which covered his earlier offerings, he began to issue catalogues that focused on items such as bulbous flowers and tubers, greenhouse plants, chrysanthemums, and vegetable and flower seeds. William Jr. attracted additional attention in 1828 when he published one of the first strictly horticultural books to come from the United States: A Short Treatise on Horticulture: Embracing Descriptions of a Great Variety of Fruit and Ornamental Trees and Shrubs, Grape Vines, Bulbous Flowers, Green-House Trees and Plants, &c. The book described all the plant offerings at the Linnaean Botanic Garden and Nursery, in some sense serving as an extended advertisement. The treatise also comprehensively covered horticultural topics, such as planting, pruning, and propagation. It even included information about soil preferences and methods for fungal disease control. Over the next three years, William Jr. worked with his son, William Robert, on two additional books, for which his son was the primary author. The first, A Treatise on the Vine, was published in 1830 and was the first significant book written in America on grape cultivation. The Princes had systematically tested scores of European grape varieties (Vitis vinifera), along with improved varieties of native North American grapes (like V. labrusca and V. riparia), and interspecific hybrids. The book described over two hundred European grape varieties and eighty American. This work helped to establish viticulture as a fullfledged branch of American horticulture, and for William Robert, grape breeding and cultivation remained a lifelong interest. The second book, The Pomological Manual, published in 1831, was a two-volume cyclopedia that attempted to catalogue all fruit varietwith two solid Dutch doors on either end and a bust of Linnaeus (likely from his father) on a bracket against the wall. The house's formal gardens contained two ginkgos (Ginkgo biloba), which were among the oldest in the country, and an old cedar of Lebanon (Cedrus libani) that the Princes had imported from France. Under William Robert's leadership, however, the business began to struggle. In the 1830s, he speculated heavily in the domestic silk industry and may have been a key contributor to the skyrocketing prices for mulberry trees (Morus alba), the food source for silkworms. He imported more than one million mulberry trees from France in 1839, and shortly afterward, the price for mulberry trees crashed. When this venture failed, the Princes could not keep up with mortgage payments on the nursery, and by 1841, they lost the Linnaean Botanic Garden and Nurseries in foreclosure. These events spawned a bitter controversy with the property's new owner, Gabriel Winter, who was married to one of William Jr.'s cousins. Although William Robert continued to raise and sell plants from an adjacent nursery property, he and Winter competed in horticultural publications over the right to sell plants as the Linnaean Botanic Garden and Nurseries. Ultimately, the Princes kept the name, and Winter sold the remaining plant inventory and subdivided the original property for housing development. By 1846, the finances at the new Prince nursery began to stabilize, and William Robert published Prince's Manual of Roses, his third and final significant contribution to horticultural literature. At his new botanic garden, William Robert grew over seven hundred rose varieties, and the book provided detailed descriptions of varieties and featured many roses from China. He also included information about horticultural care and propagation. It was one of the very best works on this subject. Still, it was eclipsed in popularity by Samuel B. Parsons's book published the following year: The Rose: Its History, Poetry, Culture, and Classification. Parsons\u2014the proprietor of Parsons Nursery in Flushing\u2014ultimately revised his book as Parsons on the Rose: A Treatise on the Propagation, Culture, and History of the Rose. The competition between these books suggests the horticultural foment that was occurring in Flushing during this period. William Prince Jr. and his son William Robert Prince (above) authored seminal American horticultural manuals. In A Treatise on the Vine, published in 1830, they promoted new grape varieties, including 'Isabella', which became a favorite of American viticulturists. HEDRICK, 1908 AND 1911\/ARCHIVE OF THE ARNOLD ARBORETUM 22 Arnoldia 78\/5-6 \u2022 October 2021 Later, William Robert went on two extended botanical expeditions, to California (in 1849) and Mexico (in 1850). While these trips suggest that the business was doing reasonably well, William Robert began to gradually withdraw from the day-to-day management of the nursery around 1855, at the age of sixty. Instead, he devoted his energy to other botanical interests, including research on botanical medicinal remedies. He also continued to breed and evaluate new varieties of fruits and ornamental plants, especially grapes, strawberries, and roses. His oldest son, William III, meanwhile assumed increasing responsibility for the enterprise. William Robert's career reflected the changes that were going on in the American horticultural community. His father had been a founding member of the New York Horticultural Society in 1818 and joined the Massachusetts Horticultural Society after it was established in 1829, but he was also a member of the Linnaean Society of Paris, the Horticultural Society of London and Paris, and the Academy of Georgofili, based in Florence, Italy. William Robert invested his energy into the increasingly sophisticated American horticultural societies rather than those in Europe. He contributed many articles to the leading American agricultural magazines of the day, such as The Rural New Yorker and Gardener's Monthly. Moreover, he was a member of the American Institute of the City of New York and the American Pomological Society. On March 28, 1869, William Robert died at his home in Flushing, and as it turned out, the esteemed business died with him. William III had enlisted for the United States Army during the Civil War, and he chose to remain in the military. William Robert's second son, LeBaron Bradford, pursued a career in law and politics. Gardener's Monthly printed a two-page obituary for William Robert. It was a sad and respectful tribute to his horticultural brilliance while nonetheless remarking on his combative personality. Meanwhile, the Massachusetts Horticultural Society issued a full resolution commemorating his life as a \"pioneer in the field of horticulture,\" a title that seems equally appropriate for the three generations of Princes that came before him. In 1939, efforts were made to move William Robert's house to the site of the New York World's Fair, to demonstrate a historic colonial homestead, but the campaign came to no avail. Later, New York City park commissioner Robert Moses rejected a proposal to move the structure to Flushing Meadow Park. Moses's vision for a \"modern city\" had little space for old wooden buildings. In its last few years of use, the structure served as a rooming house and a club. The shabby, unpainted building was then boarded up and surrounded by billboards and a gas station. The house was torn down in 1942. Of course, by that point, the lush greenhouses that once welcomed winter visitors had long ago disappeared, and the nursery property had been subdivided and sold for development. Yet, the 150-year story of the Prince family lives on today. The family built a foundation for commercial horticulture in the United States. They championed the cultivation of plants from across the country and around the world, and their publications promoted best practices in horticulture. They even helped with establishing a more systematic approach for horticultural nomenclature. Moreover, the success of the Prince nurseries is inextricably linked to the subsequent generation of horticulturists who established businesses in Flushing. This expanding group of nursery owners became leaders in their own right. In this way, a horticultural legacy that began with one family who lived on the edge of Flushing Creek became a national and international story. Acknowledgment I'm grateful for the support of Susan Lacerte, who recently retired as executive director at the Queens Botanical Garden, located near the former Prince Nurseries. Susan's knowledge of horticulture in Flushing, both present and historical, has been an inspiration. References Cornett, P. 2004, January. Encounters with America's premier nursery and botanic garden. Twinleaf: 1-12. Downing, A. J. 1845. The fruits and fruit trees of America: Or the culture, propagation and management in the garden and orchard of fruit trees generally; with descriptions of all the finest varieties of fruit, native and foreign in this country. New York: Wiley and Putnam. \u222b The Prince Family 23 Gager, C. S. 1912, October. The first botanic garden on Long Island. Brooklyn Botanic Garden Record, 1(4): 97-99. Hedrick, U. P. 1911. The plums of New York (Report of the New York Agricultural Experiment Station). Albany, N.Y.: J. B. Lyon Company. Hedrick, U. P. 1908. The grapes of New York (Report of the New York Agricultural Experiment Station). Albany, N.Y.: J. B. Lyon Company. Hedrick, U. P. 1925. The small fruits of New York (Report of the New York Agricultural Experiment Station). Albany, N.Y.: J. B. Lyon Company. Hedrick, U.P. 1933. A history of agriculture in the State of New York. Printed for the New York State Agricultural Society, Albany, N.Y.: J. B. Lyon Company. Hotchkiss, T. W. 1934. Prince house, Lawrence Street & Northern Boulevard, photographs, written historical and descriptive data. Dist. No. 4, Southern New York State, Historic American Building Survey, HABS No. 4-19. Jacobsen, A. and Williams, J. D. 2009. Prince family nurseries (ca. 1737- post- 1851). Bulletin of the Hunt Institute of Botanical Documentation, 21(1): 4-7. Johnson, J. 1887. The village of Flushing, map of desirable building lots, Flushing: A historical sketch. New York: John P. Stock, Printer. Manks, D. S. 1967. How the American nursery trade began. Plants & Gardens, 23(2). McGourty, F. 1967. Long Island's famous nurseries. Plants & Gardens, 23(3). Munsell, W. W. 1882. History of Queens County, New York, with illustrations, portraits, & sketches of prominent families and individuals. New York: Press of George MacNamara. Prince, B. and Mills, S. F. 1823. A treatise and catalogue of fruit and ornamental trees, shrubs, &c., cultivated at the Old American Nursery. New York: Wm. Grattan. Prince, W. 1771. To be sold, by William Prince, at Flushing-Landing, on Long-Island, near New- York, a large collection of fruit trees, as follows. New York: H. Gaine. Prince, W. 1790. To be sold, by William Prince, at Flushing-Landing, on Long-Island, near New- York, a large collection, as follow, of fruit trees and shrubs. New York: H. Gaine. Prince, W. 1825. Annual catalogue of fruit and ornamental trees and plants, bulbous flower roots, green-house plants, &c. &c., cultivated at the Linnaean Botanic Garden, William Prince, proprietor. New York: T. and J. Swords. Prince, W. 1828. A short treatise on horticulture: Embracing descriptions of a great variety of fruit and ornamental trees and shrubs, grape vines, bulbous flowers, green-house trees and plants. New York: T. and J. Swords. Prince, W. R. and Prince, W. 1830. A treatise on the vine; Embracing it history from the earliest ages to the present day, with descriptions of above two hundred foreign, and eighty American varieties, together with a complete dissertation of the established culture, and management of vineyards. New York: T. & J. Swords. Prince, W. R. and Prince, W. 1831. The pomological manual; or a treatise on fruits: containing descriptions of a great number of the most valuable varieties for the orchard and garden. New York: T. & J. Swords. Prince, W. R. 1846. Princes' Manual of roses: Comprising the most complete history of the rose, including every class, and all the most admirable varieties that have appeared in Europe and America, together with ample information on their culture and propagation. New York: Clark & Austen, Saxton & Miles, Wiley & Putnam, and Stanford & Swords. Ross, P. 1902. A history of Long Island: From its earliest settlement to the present time. New York: Lewis Publishing Co. Smith, E. A. and Hayward, G. 1841. The village of Flushing, Queens County, L.I.: nine miles east of the city of New York: lat. 40\u00b0 45' 1\"N, lon. 73\u00b0 09' 58\"W. [Flushing?: s.n., ?] [Map] Retrieved from the Library of Congress, https:\/\/www.loc. gov\/item\/2008620796 St. George's Episcopal Church, Baptismal Records, 1800- 1840, Flushing, N.Y., 135-32 38th Avenue, Flushing, N.Y., Rev. Wilfredo Benitez, Rector. Trebor, H. (Ed.) 1938, October. Garden center: The four Princes\u2014William of America. So This is Flushing. Flushing, N.Y.: Halleran. U.S. Department of Agriculture. 1976. The Prince family manuscript collection: A register of their papers, in the National Agricultural Library (Library list 101). Beltsville, MD: U.S. Department of Agriculture, National Agricultural Library. Waldron, R. K. 1958. Prince's plants. The Call Number, 20(1). J. Stephen Casscles is an attorney, winemaker, and horticultural writer living in the Hudson Valley. His publications include Grapes of the Hudson Valley and Other Cool Climate Regions of the United States and Canada, published by Flint Mine Press.","distinct_key":"arnoldia-2021-Such a Fine Assemblage: The Jesup Collection of North American Woods"},{"has_event_date":0,"type":"arnoldia","title":"Thomas Meehan: The Horticultural Popularizer","article_sequence":8,"start_page":50,"end_page":61,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25743","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160af6b.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Aiello, Anthony S.","article_content":"Aside from details exchanged among horticultural history buffs or students of botanical Latin (who know Meehania, a genus in the mint family), little is widely known or remembered of the life and work of Thomas Meehan, a Philadelphia nurseryman, author, editor, and social reformer who rose to prominence in the second half of the nineteenth century.1 Meehan immigrated to Philadelphia when it was still a set of disparate and unincorporated townships on the cusp of transformation into a major industrial city. Upon his arrival, he inherited a horticultural mantle from the Philadelphia Quakers who had studied the flora of the eastern United States and built notable collections of plants in their gardens. Meehan looked to these established collections and assumed the role of the horticultural popularizer. During his long career, he used his nursery and publications to encourage the cultivation of an ever-widening palette of plants. Meehan's desire to engage a broad horticultural audience was clear from the start. In his first book, The American Handbook of Ornamental Trees, published in 1853, Meehan described his intention of creating something for \"extensive popular use.\"2 This goal persisted as he continued to write and edit a series of prominent horticultural magazines, and towards the end of Meehan's career, Charles Sprague Sargent, the director of the Arnold Arboretum, described Meehan's accomplishments as \"a most important factor in increasing the cultivation of American trees and shrubs.\"3 In Philadelphia, Meehan led a remarkable life, contributing to a staggering array of fields. His work is hard to encapsulate, so this article will not offer a complete accounting; instead, to use Meehan's own words, it will present \"an anthology, and will not aim at anything further than to cull the most beautiful, interesting, and important.\"4 Thomas Meehan: The Horticultural Popularizer Anthony S. Aiello At Bartram's Garden Meehan was born in Potter's Bar near London, England, in 1826. From an early age, he was trained in horticulture by his father, himself a well-known gardener. Meehan held several prominent gardener positions as a teenager, before pursuing his formal education at the Royal Botanic Gardens, Kew, graduating in 1848.5 Having been refused a gardening position in England based on religious grounds, Meehan saw the opportunities offered in the United States. By March of the same year, he arrived in Philadelphia, where he would spend the remaining fifty-three years of his life.6 Once in Philadelphia, Meehan quickly became acquainted with the leading horticulturists of the city. He began by working for Robert Buist, who was establishing Rosedale Nursery on what was then the rural edge of southwest Philadelphia. The nursery was famous for its seed business and its selections of fruit and ornamental trees. After one year with Buist, Meehan accepted an offer to work at Bartram's Garden.7 At that point, the garden was transitioning from ownership by the Bartram family to Andrew M. Eastwick, a railroad magnate, who had recognized the garden's importance and built an elaborate Victorian home there, preserving the original Bartram house and its famous plant collection. Until 1850, Bartram's Garden had been operated by the founding family. John Bartram, the patriarch, had been a royal botanist for the king of Great Britain. He and his son William explored the eastern United States, collecting seeds that they propagated for their garden and distributed to other respected horticulturists throughout America and Great Britain. William maintained the garden upon his father's death. In turn, William's niece Ann Bartram Carr and her husband, Robert, were the third generation AIELLO, A. S. 2021. THOMAS MEEHAN: THE HORTICULTURAL POPULARIZER. ARNOLDIA, 78(5-6): 50-61 Facing page: Thomas Meehan was a central booster of American horticulture in the late nineteenth century. As a nursery owner, he promoted an ever-widening palette of plants, and as a horticultural writer and editor, he did the same. He is photographed here for the Centennial Exposition of 1876. PORTRAIT COURTESY OF THE FREE LIBRARY OF PHILADELPHIA, PRINT AND PICTURE COLLECTION Earth 51 52 Arnoldia 78\/5-6 \u2022 October 2021 to build the collection, continuing the family's international trade in seeds and plants. One can only imagine Meehan's fascination with this plant collection, undoubtedly one of the best in the United States at the time and one primed for study by a keen student of horticulture. While he was there, Meehan began collecting notes for his first book, The American Handbook of Ornamental Trees. He fitted out a place to write in the woodshed that John Bartram had used for potting and packing seed.8 It is difficult to imagine what Meehan's experience was like in that woodshed, but from a photograph that he published of the structure years later, it appears analogous to an artist's garret, cramped quarters but perhaps a place with little to distract the author from his work. In the garden, what would Meehan have experienced? From the Handbook, published in 1853, we get a sense of the diversity and size of the trees growing there. Fittingly, many of the trees that Meehan described would have been potted up in the very same building where he collected his observations as much as a century later. Meehan first intended for the book to list the trees growing at Bartram's Garden, but it evolved into a more comprehensive project that included all the trees (and some shrubs) cultivated throughout the Delaware Valley and presumably across the Northeast. In 1852, while he worked on the project, Meehan left Bartram's Garden to work for Caleb Cope, the former president of the Pennsylvania Horticultural Society. Cope's Springbrook estate was located along the Delaware River in far northern Philadelphia.9 In presenting his authorial credentials, Meehan acknowledged his time at Kew and several \"superior establishments\" in Philadelphia. He added that \"nothing has been admitted into the body of the work that has not been the result of the personal experience of the author. No tree is described as being in cultivation which the author has not himself seen.\" Meehan's horticultural ambitions are evident from his ability to visit and bear first-hand witness to so many trees in such a short period. The pace is even more remarkable given that travel on unimproved roads among the surrounding counties was challenging. Yet, Meehan's inveterate field research not only allowed him to understand the regional horticultural diversity but also brought him into the gardens of prominent botanical collectors. The Handbook documented the gardens of the early Philadelphia Quaker botanists and described the transition from the local horticultural heritage to a broader palette of plants from Europe and Asia. Here we see Meehan serving as a bridge between two eras: from the horticultural legacy of the late 1700s and early 1800s to the broader and more outward-looking horticultural developments of the late nineteenth century. The Handbook provides glimpses into the most renowned collections of the time. Of course, Meehan describes numerous notable trees at Bartram's Garden, including an old Franklin tree (Franklinia alatamaha, listed as Gordonia pubescens), which was likely one of William Bartram's original eighteenth-century collections. Meehan also lists massive specimens like a ninety-three-foot-tall Kentucky coffeetree (Gymnocladus dioicus) and a fifteenfoot- tall cornelian-cherry (Cornus mas), a European species that would have been a collector's tree at that time. Meehan also describes plants at the home of Humphry Marshall\u2014author of Arbustrum Americanum: The American Grove, who lived near West Chester\u2014and the now-forgotten arboretum of John Evans, which was one of the most significant collections of its time, located in Radnor, about fifteen miles west of Philadelphia. The best extant example of a nineteenthcentury arboretum that Meehan visited is that of the Peirce family, which now comprises the core of Peirce's Park at Longwood Gardens. The Peirces began their collection in the early 1800s, creating one of the finest regional arboreta by building on their forerunners, the Bartrams and Marshalls. The collection became renowned for its scale and diversity. Meehan describes several notable trees at this location, some of which remain today. For example, in his description of eastern redbud (Cercis canadensis), Meehan mentioned that he had \"seen fine specimens of this in Mr. Pierce's [sic] fine avenue.\" Similarly, he listed a cucumbertree magnolia (Magnolia acuminata var. subcordata, then M. cordata) with a four-foot circumference in Peirce's arboretum. In recent years, this tree was named as the cultivar 'Peirce's Park', and although the original tree was lost during a storm in April Thomas Meehan 53 2020, several young ones are planted throughout Longwood Gardens. Meehan's horticultural explorations were not limited to prestigious gardens. A favorite tree citation in the Handbook is of paper mulberry (Broussonetia papyrifera), a curious species native to East Asia. Meehan wrote that it \"thrives on the sea-shore,\" growing in Cape May, New Jersey. Boat travel from Philadelphia to Cape May was then much easier than overland travel, and Cape May's geography led to its development as a Victorian-era resort. One can picture Meehan taking a busman's holiday to the beach, recording notes even during precious personal time. At the time, he would have been courting his future wife, Catherine (Kitty) Colflesh, and one can imagine her joining him on tree-hunting excursions. Meehan's appendix is equally informative for students of horticultural history because it lists tree species recently introduced but which he had not observed. This detail helps to date the introduction of these species into the United States, or specifically Philadelphia. For example, Meehan lists nine species of maple in the main text: six native to the eastern United States, along with two common European species, the hedge maple (Acer campestre) and Norway maple (A. platanoides). In his appendix, however, he listed maples that he was aware of but had not seen. These included the vine maple (A. circinatum) from the Pacific Northwest, and the Bosnian and Italian maples (A. obtusatum, and A. opalus, respectively), which were just appearing on the East Coast. Germantown Nurseries In 1854, Meehan started a nursery in partnership with William Saunders of Baltimore in the Germantown section of Philadelphia, well outside the developed portions of the city.10 While Saunders's involvement lasted only a year, the Germantown Nurseries quickly became one of the regional leaders in growing and selling trees, shrubs, and perennials. Meehan's brother Joseph joined the operation in 1859, and his Thomas Meehan compiled notes for his first book in John Bartram's woodshed\u2014a place where the Bartram family likely potted some of the very trees that Meehan described decades later. ARCHIVES OF THE ARNOLD ARBORETUM; MEEHAN'S MONTHLY (VOL. 6) 54 Arnoldia 78\/5-6 \u2022 October 2021 three sons (Thomas B., J. Franklin, and S. Mendelson) came on board in the decades to come. As evidence of the success of the operations, what had begun as a few acres of land in Germantown grew to 75 acres by the late 1800s and then to 150 acres by the turn of the twentieth century, encompassing property in Germantown and suburban Dresher, Pennsylvania.11 The nursery was especially known for its diverse offerings of North American trees. By 1893, a correspondent for Garden and Forest noted that \"Mr. Meehan early recognized that \u2026 American plants are the best for America\" and went on to say that \"in no other place are American trees and shrubs raised in such quantities.\" Their offerings included native species that were difficult to find at other nurseries. Yet, Meehan simultaneously offered and promoted non-natives species as they became available.12 This Janus-like approach to horticulture continued the link to Philadelphia's horticultural heritage while recognizing the changing demography and tastes of the city's gardeners. American nursery catalogues from the mid- 1800s reveal that most ornamental trees offered were from North America and Europe, with a smattering from Asia Minor and Asia.13 A watershed moment in the availability of greater plant diversity occurred at the Centennial Exposition, the first official world's fair held in the United States, which took place in Philadelphia from the spring to autumn of 1876. As a celebration of the one-hundredth anniversary of the signing of the Declaration of Independence, the event exposed a vast audience to a wide array of modern conveniences, inventions, and international cultures. Also, through various horticultural exhibits, the Exposition introduced Asian (particularly Japanese) plant species to a broad American audience. Prior to the Exposition, GERMANTOWN HISTORICAL SOCIETY\/HISTORIC GERMANTOWN Elms flank the entrance to Meehan's Nurseries, photographed around 1902. Thomas Meehan 55 Japanese species were slowly making their way into Boston and New York but had yet to see wider availability.14 Meehan created an arboretum of over seven hundred trees for the Exposition. Local newspapers described it as a \"grand miniature forest\" that was especially noteworthy for its collection of \"trees and shrubs of the United States.\"15 Other prominent nurserymen had displays nearby, including Josiah Hoopes (whose display included twelve hundred evergreens and forty varieties of ivies), Robert Buist (showcasing trees, shrubs, and herbaceous plants), and S. B. Parson & Sons (who were reported to have \"remarkable Japanese plants, maples, evergreens, azalias [sic], new shrubs, and half hardy plants\").16 After the Exhibition, Meehan and the other nursery owners provided portions of their outdoor collections to Philadelphia's Fairmount Park. Therefore, the diversity of their displays is suggested in Joseph Rothrock's catalogue of the trees and shrubs in Fairmount Park, published in 1880. The catalogue documents early introductions of Asian species, including Japanese maple (Acer palmatum), Asian magnolias (like Magnolia campbellii and M. denudata), panicle hydrangea (Hydrangea paniculata), and the lacebark pine (Pinus bungeana).17 After the event, the diversity of plant offerings from Japan rapidly increased, and by the end of the 1800s, many now-familiar plants, and many that we still think of as \"rare and unusual,\" were regularly offered for sale. Meehan was quick to recognize the importance of these introductions. When he wrote about the other nursery displays at the Exhibition in Gardener's Monthly, a magazine that he had edited since 1859, he remarked on the \"special bed\" of Japanese plants shown by S. B. Parsons & Sons. Among the most striking plants, he reported, was the red-leaved Japanese maple (now Acer palmatum forma atropurpureum).18 By 1882, Meehan's nursery catalogue offered one-foot-tall specimens of this for two dollars, then among his most expensive offerings. On the back cover of the same catalogue, he proudly advertised the \"Japan Snowball\" (Viburnum plicatum), claiming that his nursery had been first to introduce it into the United States. This claim was accompanied by the only illustration in the catalogue, suggesting that Meehan fully recognized the commercial importance of these newcomers.19 By the 1890s, Meehan's nurseries were offering a weeping Japanese cherry (what would now be considered Prunus subhirtella), Asian magnolias and maples, and even umbrella pine (Sciadopitys verticillata) and Hiba falsearborvitae (Thujopsis dolabrata).20 In some sense, Meehan's nursery served as a laboratory for him to study plants. A perfect example of this is the daimyo oak (Quercus dentata). At a meeting of the Academy of Natural Sciences of Philadelphia in 1886, Meehan presented a short description of the floral structure of Quercus dentata, grown from seed that he had received from Japan at the time of the Centennial Exposition.21 By 1895, the daimyo oak was offered by his nursery, described as \"a rich addition to our list of oaks \u2026 in May the yellow flowers, in long aments, make it attractive in a way no other oak is.\"22 Despite his ever-increasing interest in nonnative species, Meehan maintained a strong affinity for native plants. In the same 1895 catalogue in which he advertised the daimyo oak, Meehan wrote that \"for twenty years or more we have been trying to impress upon American planters the importance of using Native Oaks in landscape works \u2026 and finally, after all these years, planters began to realize that we were right and to recognize in the American Oak, the 'King of Trees.'\"23 And while Meehan is often most associated with woody plants, his catalogues have a large diversity of native herbaceous perennials and hardy ferns\u2014many sought out by today's keen gardeners. Meehan's nursery distributed plants to botanical institutions, including the Arnold Arboretum where a few dozen specimens are still alive. The most historically significant are two Franklin trees (Franklinia alatamaha, accession 2428-3*A and *B), propagated in 1905 from a plant that Meehan provided about thirty years earlier. These are believed to be the oldest living representatives of the species.24 Other Meehan plants at the Arboretum include a group of five black oaks (Quercus velutina, accession 1237), acquired in 1873, when the Arboretum was only a year old, and a Southern red oak (Q. falcata, accession 3333*A). These North American oaks are now living reminders of Meehan's commitment to the \"King of Trees.\" 56 Arnoldia 78\/5-6 \u2022 October 2021 Horticultural Writer and Editor Meehan was a prolific author throughout his career. He served as editor of the Gardener's Monthly until 1888, when its publisher, Charles Marot, died. A few years later, Meehan's Monthly was born and continued until 1902. Over his forty years as the editor of monthly publications, Meehan generated a vast amount of material to read. His prodigious output is hard to encapsulate or even anthologize. The tone of the publications was conversational and newsy, and his personal writing style was both informative and approachable. In a period before easy (not to mention instant) communication, these journals regularly shared information and current trends, mixed with a bit of human interest.25 In the initial issue of Garden and Forest, in 1888, an unsigned editorial (perhaps written by Charles Sargent, who \"conducted\" the magazine) commented on the loss of the Gardener's Monthly: \"Ever since we have been interested in the cultivation of flowers we have looked to the Monthly for inspiration and advice, and its pages have rarely been turned without finding the assistance we stood in need of.\" The editorial continued by celebrating Meehan's imprint on the publication. \"Fortunately, the Gardener's Monthly, and its modest and accomplished editor, Mr. Thomas Meehan, were one and the same thing. It is Mr. Meehan's long editorial experience, high character, great learning and varied practical knowledge, which made the Gardener's Monthly what it was. These, we are happy to know, are not lost to us, as Mr. Meehan will \u2026 continue to delight and instruct the horticultural public.\"26 In the late 1870s, Meehan had also begun a multivolume work titled The Native Flowers and Ferns of the United States. The project is another testament to his long-standing love of North American plants. In the preface to the first volume, Meehan described how the project emerged from his desire to write a scientific treatment on the North American flora. Although he pitched this idea to a publisher, he ultimately decided, once again, to focus on engaging a more general audience. \"A purely scientific and systematic treatise \u2026 must necessarily be limited to a small circle of readers,\" he explained, \"and even in this small circle there would be but a few who would care to subscribe to a work, the end of which they might never live to see.\" Four volumes were produced, and Meehan's voice shines through them. He lushly described almost fifty species in each volume, often incorporating history, poetry, and horticultural information. The entry for each species included a lavish color illustration.27 The project was revived in 1891 when Meehan's Monthly was launched. While Meehan's Monthly was a newsy horticultural periodical, in keeping with the style and tone of the Gardener's Monthly, each issue began with a description of a native species and was accompanied with illustrations prepared for unpublished volumes of the Native Flowers and Ferns project. Garden and Forest celebrated the arrival of this new periodical: \"Mr. Meehan's return to horticultural journalism will be welcomed by many readers of the Gardeners' Monthly who felt something like a personal bereavement at the discontinuance of that excellent magazine.\"28 Along with these horticultural pursuits, Meehan maintained a long-running correspondence with many notable botanists of his time, including George Engelmann, Asa Gray, and Charles Darwin. Much of this correspondence concerns specific observations or botanical questions, often relating to articles that Meehan would eventually publish in the Proceedings of the Academy of Natural Sciences of Philadelphia, where he long served as the vice president. Advocate for Urban Green Space In the later years of his life, Meehan became actively involved in urban improvement. In 1883, he accepted a role on the Philadelphia Common Council in order to ensure the creation of city parks and preservation of Bartram's Garden.29 Meehan was instrumental in forming the City Parks Association, creating lasting green space in the most urbanized neighborhoods. He is credited with introducing nature study and kindergarten to Philadelphia public schools, and he strived to improve the educational system for working-class families throughout the city.30 Among these accomplishments, it is the preservation of Bartram's Garden that is the most noteworthy. In 1879, Andrew Eastwick died, Thomas Meehan 57 Thomas Meehan's work on The Native Flowers and Ferns of the United States serves as one of the clearest examples of his lucid writing style. Each of his entries was accompanied by chromolithograph illustrations prepared by Louis Prang of Boston. The illustrations and excerpts here appeared in later installments of the project in Meehan's Monthly. Pinkshell Azalea (Rhododendron vaseyi) \" It is one of a number of beautiful plants missed by the early explorers of the Mountains of North Carolina, and which have been brought to light only in modern times.\" MEEHAN'S MONTHLY (VOL. 7) ILLUSTRATIONS FROM THE ARCHIVES OF THE ARNOLD ARBORETUM 58 Arnoldia 78\/5-6 \u2022 October 2021 Coast Cholla (Cylindropuntia prolifera) \" Animals take the fruit to their haunts, use the fl esh, and scatter the undigested seeds in various directions,\u2014certainly many fruit-bearing plants are widely distributed in this manner. Those who think this feature a special adaptation will see in the absence of spines in the fruit of this species, strong confi rmation of this view. The plant would be spiny, it would be contended, in order to protect it against browsing creatures; while, when consumption instead of protection became useful to the plant, the production of spines would be arrested.\" MEEHAN'S MONTHLY (VOL. 3) Thomas Meehan 59 Rosebay Rhododendron (Rhododendron maximum) \" In a state of nature the Rhododendron inhabits wild, rocky places, in uninhabited regions where the foot of the traveler is rarely seen \u2026 So far away are they generally in their gloomy homes that even the great traveler, John Bartram, had not met with them anywhere west of the Schuylkill river.\" MEEHAN'S MONTHLY (VOL. 1) 60 Arnoldia 78\/5-6 \u2022 October 2021 Partridgeberry (Mitchella repens) \" It is remarkable that a plant so attractive in so many ways should not have become more attached to the public mind, or received more attention from polite writers, but the author can recall no instance in American poetry or general literature in which the Partridge berry plays a conspicuous part.\" MEEHAN'S MONTHLY (VOL. 3) and for nearly a decade, the resolution of his estate and the fate of Bartram's Garden remained unresolved.31 Shortly after Eastwick's death, Sargent, using his connections in Philadelphia, tried to organize a group of \"liberal gentlemen\" to purchase the property.32 This effort was unsuccessful because the owners of the estate believed that \"they could make more [profi t] by destroying its botanical associations, and turning the whole into building lots.\"33 Sargent continued to provide support on a national level through Garden and Forest, arguing in an unsigned editorial that \"the name of Bartram's Garden should be preserved and \u2026 should be maintained in as near the condition as its fi rst owner left it.\"34 Meanwhile, Meehan and members of the City Parks Association continued the local campaign. Ultimately, the City of Philadelphia appropriated funds to purchase Bartram's Garden in 1889, took ownership in 1891, and fi nalized the purchase in 1893.35 As a result, more than forty years after Meehan had fi rst worked at the historic garden, it became preserved in perpetuity. This achievement must have been remarkably gratifying for Meehan, seeing the preservation of the place that helped to launch his career and that had such horticultural signifi cance in his adopted city. Once the future of Bartram's Garden was settled, Meehan's foresight in creating open space throughout the city was acknowledged with another Garden and Forest editorial: \"The fact that the people of Philadelphia are securing a series of small parks is largely due to the publicspirited and tireless efforts of Mr. Thomas Meehan, the well-known horticulturist \u2026 Many generations of Philadelphians will have a good reason to remember with gratitude his disinterested efforts for the improvement and happiness of his fellow men.\"36 Meehan's Legacy As a coda to his life, Meehan was awarded the Veitch Memorial Medal in 1901, a few months before he died. He followed Sargent and Liberty Hyde Bailey as the third American to win this honor. In conferring it, the Royal Horticultural Society recognized his \"distinguished services in botany and horticulture.\" Seeing Meehan in the company of these two towering fi gures of late nineteenth and early twentieth-century Thomas Meehan 61 American horticulture affirms his stature among his peers: Sargent, one of the great dendrologists of his era, who brought the Arnold Arboretum to prominence, and Bailey, a man of astoundingly broad interests and accomplishments who combined the science of botany with the art of horticulture. Meehan pursued similar combinations and was interested not only in the world of horticulture but in using it for the betterment of his fellow citizens. It is worth pondering what Meehan would think if he were to see the state of contemporary horticulture. Certainly, many if not most of the trees that are commonly planted across the Northeast would be familiar to him. Having straddled the divide between native and nonnative plants, he might think that there would be no need for invidious comparisons between the two groups. And he might be bemused at the trends in \"new\" native plants, having promoted many of those species in his various publications and through his nursery. If nothing else, although his name may have faded, Thomas Meehan's impact as a promoter of modern horticulture has not. Endnotes 1 Oberle, S. G. 1997. The influence of Thomas Meehan on horticulture in the United States. University of Delaware, M. S. Thesis Dissertation 2 Meehan, T. 1853. The American handbook of ornamental trees. Philadelphia: Lippincott, Grambo, and Co. 3 Sargent, C. S. 1890. Silva of North America (vol. 9). New York: Peter Smith. 4 Meehan, T. 1878. The native flowers and ferns of the United States in their botanical, horticultural, and popular aspects (vol. 1). Boston: L. Prang. 5 Meehan, S. M. 1902. A brief sketch of the life of Thomas Meehan. Meehan's Monthly, 12: 13-19. 6 Harshberger, J. W. 1899. The botanists of Philadelphia and their work. Philadelphia: T. C. Davis. 7 Meehan, T. 1896a. Meehan letter to C. S. Sargent, 16 August 1896. Charles Sprague Sargent (1841-1927) papers, Arnold Arboretum Horticultural Library, Harvard University. 8 Meehan, T. 1896b. John Bartram's wood-shed. Meehan's Monthly, 6: 17. 9 Meehan, 1902. 10 Meehan, 1902. 11 Oberle, 1997. 12 S. 1893, September. The Meehan Nurseries and the trees of Germantown. Garden and Forest, 6(289): 377-378. 13 See, for instance: Hoopes, Bro. & Thomas. 1870. Annual trade list of the Cherry Hill Nurseries, West Chester, Pa.: Spring of 1870. West Chester, PA: Hoopes, Bro. & Thomas. 14 Del Tredici, P. 2017. The introduction of Japanese plants into North America. The Botanical Review, 83: 215-252. 15 Thomas Meehan of Germantown. 1876, April. Reading Times (Reading, PA), 37(22): 2; An interesting display. 1876, May. The Daily Evening Express (Lancaster, PA), 39(105): 2. 16 Burr, S. J. 1877. Memorial of the International exhibition. Hartford: L. Stebbins. 17 Rothrock, J. T. 1880. Catalogue of trees and shrubs native of and introduced in the horticultural gardens adjacent to Horticultural Hall in Fairmount Park, Philadelphia. 18 Meehan, T. 1876. Horticulture at the Centennial. The Gardener's Monthly, 18(212): 254-256. 19 Germantown Nurseries. 1882. General price list for the fall of 1882. Germantown, PA: Germantown Nurseries. 20 Meehans' Nurseries. 1895. Catalogue. Germantown, PA: Thomas Meehan & Sons. 21 Meehan, T. 1886. Note on Quercus dentata. Proceedings of the Academy of Natural Sciences of Philadelphia. 38: 280-281. 22 Meehans' Nurseries, 1895. 23 Meehans' Nurseries, 1895. 24 Del Tredici, P. 2005. Against all odds: Growing Franklinia in Boston. Arnoldia, 63(4): 2-7. 25 Oberle, 1997. 26 The Gardener's Monthly. 1888, February. Garden and Forest, (1)1: 1. 27 Meehan, 1878. 28 Notes. 1891, March. Garden and Forest, 4(161): 144. 29 Meehan, T. 1897. In Bartram's Garden. Meehan's Monthly, 7: 50. 30 Harshberger, 1899; Meehan, 1902. 31 Fry, J. 2004. John Bartram House and Garden. Historic American Landscape Survey, (HALS) PA-1. 32 Fry, 2004; Meehan, T. 1885. The old botanic garden of Bartram. The Gardener's Monthly and Horticulturist, 27: 26-27. 33 Meehan, 1885. 34 Notes. 1889, February. Garden and Forest, 2(52): 86. 35 Fry, 2004. 36 Notes. 1889, March. Garden and Forest, 2(54): 120. Anthony S. Aiello is the associate director of conservation, plant breeding, and collections at Longwood Gardens."},{"has_event_date":0,"type":"arnoldia","title":"The Intertwined Attractions of Plants, Moths, and People","article_sequence":9,"start_page":62,"end_page":67,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25744","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160b36e.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Bawa, Kamaljit S.","article_content":"The Intertwined Attractions of Plants, Moths, and People Kamaljit S. Bawa It was a warm and humid night in September of 2003. In a tropical forest by the coast of Madagascar, Phil Devries, an entomologist and noted nature videographer, swatted mosquitoes hovering around his face. He had been waiting eagerly for a visitor since seven o'clock. As the night transitioned to early morning, without any signs of the visitor, the tension and anxiety in Phil's mind increased. For the visitor, Phil Devries was inconsequential; the desired object was Darwin's orchid near which Phil (or the Butterfly Man, as he is popularly known) had parked himself to photograph the orchid's pollinator. \"Good Heavens what insect can suck it,\" Charles Darwin is said to have remarked in reference to the nectar in the long floral tube of Angraecum sesquipedale, now known as the Darwin's orchid, native of Madagascar.1 Darwin had received the orchid on January 25, 1862, from James Bateman, a businessman, collector of plants, and horticulturist, who grew orchids. Darwin then famously predicted that A. sesquipedale must be pollinated by a hawkmoth with a proboscis that measured at least eleven inches in length.2 In 1903, almost forty years after Darwin intuited its existence, a hawkmoth with long mouth parts was described by Walter Rothschild and Karl Jordan. It was isolated from moth specimens collected on an earlier expedition to Madagascar by Jules Paul Mabille, a French naturalist. Rothschild and Jordan named the species Xanthopan morganii. However, it was not until 1992, a good ninety years later, that Lutz Wasserthal, a German biologist, observed X. morganii visiting the flowers of A. sesquipedale in real life. Only then was the connection between orchid flowers and moths finally confirmed.3 Visits of moths to flowers in the wild are hard to observe. And so, Wasserthal had to use large flight tents to photograph the two partners engaged in the mutually beneficial relationship. Finally, in 2003, after spending several nights in the Madagascar forest, Phil Devries was able to photograph the evasive moths visiting the flowers of A. sesquipedale in the wild\u2014at around three o'clock in the morning.4 The correlation between the length of the floral tube and the length of moth's proboscis led Darwin to infer the process of coevolution, in which natural selection favors reciprocal increases in the length of the floral tube and moth's proboscis. Heritable variation\u2014in this case, variation in floral tube and the length of proboscis in moths\u2014is the raw material on which natural selection acts. Between Darwin's original prediction and the eye-witness observation, 130 years had passed. Nothing in science comes easy. Not even for Darwin. It was Gregor Mendel, an Austrian monk, who proposed the principles of inheritance in 1865, based on his experiments with peas. From Darwin's orchids to Mendel's peas, plants have played an important role in the study of evolution. Curiously and coincidentally, both Darwin and Mendel were contemporaries, and although Mendel's work filled a critical gap in Darwin's theory of evolution by natural selection, the two men did not know of each other's work! While Darwin is noted for his work on evolution, he is much less known as an ardent botanist. He was greatly interested in the reproduction of plants, particularly orchids. He wrote several books on plants: The Power of Movement in Plants, On the Various Contrivances by Which British and Foreign Orchids Are Fertilised by Insects, On the Good Effects of Intercrossing, The Different Forms of Flowers on Plants of the Same Species, and Insectivorous Plants. Plants were critical to the formulation of his ideas both about inherent variation and how natural selection acts on this variation to enable evolution. Facing page: Darwin's orchid (Angraecum sesquipedale) is one of thousands of night-flowering plants pollinated by moths. In this case, only one pollinator can accomplish the task\u2014Xanthopan morganii. SENCKENBERG COLLECTION \/ PHOTO: SAMMLUNGSFOTOGRAFEN.DE BAWA, K. S. 2021. THE INTERTWINED ATTRACTIONS OF PLANTS, MOTHS, AND PEOPLE. ARNOLDIA, 78(5-6): 62-67 Moths and Sex Pheromones It is March 1974, and I am waiting, at evening time, under a large Luehea speciosa. The tree stands in a dry tropical forest in Guanacaste Province, Costa Rica. The previous day, I had seen its large white flowers start to bloom around eight o'clock in the evening. And so, the next day, under the tree and in the light of the moon, I staked a vantage point and started my watch. At exactly a quarter to eight, and almost like magic, the large white petals start to unfurl. In a quarter of an hour, almost a hundred flowers in my field of view have opened in near-perfect synchronicity. In my five decades of fieldwork in Costa Rica, that night was one of the most memorable and remains permanently etched in my memory. Plants depend on a wide variety of animals to get cross-pollinated. The diversity of these pollination systems is on full display in tropical evergreen forests, the world's most speciesrich ecological communities. On any given day, at any time during a short walk through the forest, one can encounter flowers of many sizes, shapes, and colors that are pollinated by insects\u2014largely bees, butterflies, and flies\u2014 and, at times, birds. For a different set of plant species that start to open their flowers around dusk and at night, insects (beetles and moths) and mammals (primarily bats) take over the role of major pollinators. All across the globe, but mostly in the tropics, tens of thousands of plant species are pollinated by an equally large number of moth species at night. Moth-pollinated flowers are almost always white and tubular, with nectar at the base of the tube. They blossom in the evening, soon after dusk, and the blooms last for one or two nights. During this time, the moths visit them frequently, making multiple forays throughout the hours of the night. Insect pollinators visit flowers for food, but, to them, flowers are more than a food source. They are also sites of mating and, often, a source of compounds that play an important role in facilitating these sexual encounters. Flowers produce a variety of volatile compounds to attract insects, such as moths. Smell plays an important role in attracting insects from afar, especially at night, when visual cues can only function once the pollinators approach the flower closely. Female moths use volatile compounds produced by flowers to synthesize sex pheromones, which they release to attract males. In some cases, the volatiles associated with the floral smell simply induce female moths to produce large amounts of sex pheromones, but in others, the female moths can absorb or ingest the volatiles and convert the compounds directly into pheromones. The males are not left behind. In some species of moths, males sequester pyrrolizidine alkaloids from flowers to use them as precursors for the synthesis of pheromones. Sometimes, the males even transfer the alkaloids to the female during mating, for the defense of eggs against predators.5 Thus, flowers play a critical role not only in the provision of food and nutrition but also in the mating and reproduction of pollinators. Evening Fragrances and Romantic Nights Thirty years later, I am in Bangalore, the techno-hub of South India. It is again late evening, and I am passing through a small market buzzing with people. Walking in front of vegetable and food stores, I am overpowered with fragrances emanating from buds and flowers of jasmine (Jasminum) strung together for hair adornments. And indeed, I see many women walking around with their long hair arranged in many different styles and adorned with strings of fragrant jasmine. Throughout remembered history, and for millennia, flowers have been a part of daily life in India, as adornments for gods and humans. The Hindu epic Ra\u02c9ma\u02c9 yana about the life of Ram, one of the most celebrated gods of Hindus, includes references to Sita, Ram's wife, decorating her hair with floral arrangements. And in a well-known epic poem written in the fourth century CE, the playwright Ka\u02c9 lida\u02c9 sa included a verse in which sensuality and pollination merge: Sensuous women in summer love weave flower earrings from fragile petals of mimosa 64 Arnoldia 78\/5-6 \u2022 October 2021 while wild bees kiss them gently 6 Anthologies of classical Tamil, written between 100 BCE and 250 CE, describe the flowers that women bear as those of jasmine. For men, too, flowers have been a bedtime adornment for ages, and the exchange of flowers between individuals has always carried unspoken and covert sexual connotations.7 From trees in Costa Rica that use flowers to attract moths to women in India who use flowers for adornment, the fundamental motives of life are the same irrespective of geographies, gender, or species. But the enchantment of union does not stop there. The collision of these seemingly different worlds gets closer and more intimate. Jasminium sumbac and other species of Jasminum are native to South India and other parts of tropical Asia. Jasmine flowers are highly fragrant, pollinated by moths, and here, too, the maximum production of aromatic compounds is between seven and eight o'clock in the evening!8 Moon and Sex Back in Costa Rica and on another moonlit night, I am driving to my campsite after a full day of fieldwork in the dry deciduous forest. Chains of white jasmine (Jasminum) are worn as a hair adornment in Tamil Nadu, India. The flowers become increasingly fragrant in the evening. MCKAY SAVAGE (CC BY 2.0) Intertwined Lives 65 There is little traffic on the Pan-American Highway, which means that I can easily observe the star-studded trees of Bombacopsis quinata, a relative of the silk cotton tree, on both sides of the road. Under the full moon, it is a beautiful sight, with a tree coming into view every few minutes. The \"stars,\" indeed, are large, white, moth-pollinated flowers, perched high in the leafless crowns of these very large trees. For the past several evenings, I have been passing by these trees in flower, but this time, the number of flowers on the trees appears to be unusually large. Flowers in this species last for a single night, but individual trees flower over many weeks, with a new batch opening every night. It seemed that the intensity of flowering was associated with lunar cycles, with the largest number of flowers opening on nights with the full moon. While, on this evening drive, I cannot confirm the correlation between the intensity of flowering and phases of the moon, researchers would later document such trends for other species. Moths are known to be more active on moonlit nights, and pollination can be more intense during a full moon for moth-pollinated species, as, for example, in Ephedra foeminea, a gymnosperm. In contrast to most gymnosperms, which are wind-pollinated, this species attracts moths by secreting a pollination drop from its cones. Individual plants produce their maximum amount of pollination drops during full moons. Meanwhile, a related species of Ephedra is wind-pollinated, and in that case, there is no connection between pollination and lunar cycles.9 Is there a general correlation between lunar cycles and pollination intensity for the thou- Flowers of Bombacopsis quinata open at sundown, seemingly more abundant in the treetops when the moon is full. REINALDO AGUILAR (CC BY-NC-SA 2.0) 66 Arnoldia 78\/5-6 \u2022 October 2021 sands of night-blooming plant species? We do not know. Recently, researchers have shown that a desert cactus (Cereus peruvianus), presumably pollinated by bats, puts on its largest display of flowers around the full moon. The species flowers over a few months with the number of flowers going up and down with the lunar cycles.10 The moon has always been associated with romance in our own human cultures. Surprisingly, there is insufficient data to establish a link between sexual activity with lunar cycles. Interestingly, though, research has shown that a larger proportion of females demonstrate ovulation during the full moon, and all genders experience higher aggression levels and less sleep.11 Intertwined in the Web of Life It is evening again, and the sex lives of plants, moths, and humans intertwine. All of these organisms use the same compounds to attract mates: smell is a main stimulant for each. Plants, indeed, cannot smell, yet floral volatiles are a major incentive for moths to visit flowers. Among the three partners, plants reign supreme. They seem to dictate the terms of the relationships. Moths, in fact, are held in bondage. They cannot attract mates without pheromones for which the plants hold the precursors. Humans also seem to be dependent on plants as intermediaries, although they, of course, can do without them. For those who study life on earth, the interconnections among plants, moths, and humans are not surprising. We are a part of the web of life that has celestial connections with other planets. These connections are vital for maintaining all lives, especially ours. We should celebrate and value these connections that enrich our lives by ceasing our assault on nature. Endnotes 1 Arditti, J., Elliott, J., Kitching I. J., and Wasserthal, L. T. 2012. \"Good Heavens what insect can suck it\"\u2014Charles Darwin, Angraecum sesquipedale and Xanthopan morganii praedicta. Botanical Journal of the Linnean Society, 169: 403-432. https:\/\/doi. org\/10.1111\/j.1095-8339.2012.01250.x 2 Netz, C. and Renner, S. S. 2017. Long-spurred Angraecum orchids and long-tongued sphingid moths on Madagascar: A time frame for Darwin's predicted Xanthopan\/Angraecum coevolution. Biological Journal of the Linnean Society, 122(2): 469-478. https:\/\/doi.org\/10.1093\/biolinnean\/blx086 3 Wasserthal, L. T. 1997. The pollinators of the Malagasy star orchids Angraecum sesquipedale, A. sororium and A. compactum and the evolution of extremely long spurs by pollinator shift. Botanica Acta, 110(5): 343-359. https:\/\/doi.org\/10.1111\/j.1438-8677.1997. tb00650.x 4 See video in: Tartaglia, E. 2015. Year of the Sphingidae\u2014Co-evolution. National Moth Week. https:\/\/nationalmothweek.org\/2015\/07\/17\/year-ofthe- sphingidae-co-evolution\/ 5 St\u00f6kl, J. and Steiger, S. 2017. Evolutionary origin of insect pheromones. Current Opinion in Insect Science, 24: 36-42. https:\/\/doi.org\/10.1016\/j.cois.2017.09.004 6 Miller, B. S. 1984. Theater of memory: The plays of Ka\u02c9 lida\u02c9 sa. New York: Columbia University Press. 7 Goody, J. 1993. The culture of flowers. Cambridge: Cambridge University Press, p. 323-324. 8 Braun, N. A. and Sim, S. 2012. Jasminum sambac flower absolutes from India and China\u2014Geographic variations. Natural Product Communications, 7(5): 645-650. https:\/\/doi.org\/10.1177\/1934578x1200700526 9 Rydin, C. and Bolinder, K. 2015. Moonlight pollination in the gymnosperm Ephedra (Gnetales). Biology Letters, 11(4): 10-13. https:\/\/doi.org\/10.1098\/ rsbl.2014.0993 10 Ben-Attia, M., Reinberg, A., Smolensky, M. H., Gadacha, W., Khedaier, A., Sani, M.,\u2026 Boughamni, N. G. 2016. Blooming rhythms of cactus Cereus peruvianus with nocturnal peak at full moon during seasons of prolonged daytime photoperiod. Chronobiology International, 33(4): 419-430. https:\/\/ doi.org\/10.3109\/07420528.2016.1157082 11 Moore, B. 2019. The effect of the lunar cycle on the female reproductive system. South Carolina Junior Academy of Science. https:\/\/scholarexchange.furman. edu\/scjas\/2019\/all\/242\/ Acknowledgments I thank my wife, Tshering Bawa, for encouraging me to write this manuscript when I first discussed the idea with her almost twenty-five years ago. A series of discussions with Rohini Nilekani about Brahma Kamal (Epiphyllum oxypetalum), a nocturnal blooming cactus from Mexico and South America, but widely naturalized in Asia, was another source of inspiration. Meena Narayanswamy suggested several improvements in the manuscript. Kamaljit S. Bawa is president of the Ashoka Trust for Research in Ecology and the Environment (ATREE), Bengaluru, India, and distinguished professor emeritus at the University of Massachusetts, Boston. Intertwined Lives 67"},{"has_event_date":0,"type":"arnoldia","title":"Otherworldly Wingnuts: Pterocarya x rehderiana","article_sequence":10,"start_page":68,"end_page":69,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25745","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160b726.jpg","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null,"authors":"Rubinstein, Jared","article_content":"A rarely visited corner at the Arnold Arboretum is nestled beneath the tall stone wall that separates the hickory collection from traffic on Centre Street. In late summer, the area feels otherworldly. The heavy overstory filters the light and cools the air; the humidity seems to increase; and densely planted shrubs block out the surrounding views and noises. The corner is dominated by a planting of seemingly colossal hybrid wingnuts (Pterocarya x rehderiana), with their drooping Spanish moss-like fruits and twisted forms. Standing next to their large multistemmed trunks can make you feel miniature. Wingnuts are closely related to hickories (Carya) and walnuts (Juglans). There are six species of Pterocarya, with native ranges clustered in China, Japan, Southeast Asia, and the Caucuses. In addition to cultivating representatives of five of the six species, the Arnold Arboretum has eight specimens of this unusual hybrid, all of which grow in this out-of-the-way corner. The oldest of the eight originated at the Arboretum from seed sent, in 1879, by Pierre Alphonse Lavall\u00e9e of the Arboretum de Segrez, outside of Paris. At the time, the Arboretum de Segrez was one of the largest in the world (and a noteworthy landscape where Marcel Proust once suffered an asthma attack but still managed to write a poem about its beauty). Lavall\u00e9e collected the seeds from a Chinese wingnut (P. stenoptera) in his arboretum, and, once they germinated in Boston, the seedlings were planted along Centre Street. Two decades later, Alfred Rehder, an Arnold Arboretum taxonomist, noticed that the trees didn't look quite like the Chinese wingnut. \"The trees in the Arnold, known as Pterocarya stenoptera \u2026 I can no longer consider, after much study, as the real species of that name,\" Rehder wrote to the German Dendrological Society in 1903, \"but now consider [them] a cross between this and P. fraxinifolia [the Caucasian wingnut], which in its characteristics almost exactly stops between the two species.\" Rehder hypothesized that pollen from a Caucasian wingnut growing at the Arboretum de Segrez must have landed on the flowers of a Chinese wingnut growing nearby. We don't know who collected and brought the Chinese and Caucasian wingnuts to Paris, but it may well have been the first time that the two species, normally separated by the thousands of miles between the Caucasus Mountains and eastern China, were growing in the same place. Rehder conferred with Camillo Schneider, a taxonomist working at the Vienna Natural History Museum, who agreed with Rehder's assessment. Based on their correspondence, Schneider published the first botanical description of the new hybrid in 1906. Writing in German in the Illustriertes Handbuch der Laubholzkunde, he identified the unique characteristics of the buds and rachises of the \"Bastardes\" growing at the Arnold Arboretum and officially named the hybrid for his friend, choosing the Latin name Pterocarya x rehderiana. Four trees (accession 1191) from Lavall\u00e9e's 1879 shipment still grow along the Centre Street wall, hidden behind the hickory collection. In addition, four neighboring trees (23119) were accessioned as seedlings from the original trees. When the wingnuts fruit in midsummer, they offer a dazzling display of long, pendulous clusters of winged nutlets (hence the common name) that dangle from what seems like every branch. One particularly large specimen, accession 1191*E, has an incredible form, with leaders that shoot up more than 125 feet and droop over the Works Progress Administrationconstructed bus shelter on Centre Street. As with many hybrids, Pterocarya x rehderiana seems to display hybrid vigor and, according to Rehder, are \"much hardier and more satisfactory than their supposed parents.\" A windstorm in October 2020 took out one of the leaders from accession 1191*E, but overall, the hybrids don't seem terribly affected by the cold New England winter, even after more than 140 years growing at the Arboretum. While the hybrids are a product of a chance cross that would likely have never been possible in the wild, the trees have more than claimed their uncanny home. Jared Rubinstein is an associate project manager at the Arnold Arboretum. For more on the taxonomic history of the Rehder wingnut, see his 2020 article with Michael Dosmann in Novon, issue 28(4). Otherworldly Wingnuts: Pterocarya x rehderiana Jared Rubinstein RUBINSTEIN, J. 2021. OTHERWORLDLY WINGNUTS: PTEROCARYA x REHDERIANA. ARNOLDIA, 78(5-6): 68-69"},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25699","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270af27.jpg","title":"2021-78-5-6","volume":78,"issue_number":"5-6","year":2021,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Field Botany in the Time of COVID-19","article_sequence":1,"start_page":2,"end_page":6,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25729","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25eb328.jpg","volume":78,"issue_number":4,"year":2021,"series":null,"season":null,"authors":"Brown, Emma; Maynard, Brian","article_content":"The students in our University of Rhode Island field botany class exclaimed with surprise as they tried to balance atop lopsided hummocks of tussock sedge (Carex stricta). The mounds arose between expanses of boot-sucking sphagnum moss. Red cranberries (Vaccinium macrocarpon) dotted the shimmering surface around them. This was the first time most of the students had seen cranberries in the wild\u2014a powerful learning moment. Memories of the sour explosion of the cranberries would become associated with the comradery of learning how to differentiate this flowing fen from a typical bog\u2014or how to identify the three-way sedge (Dulichium arundinaceum) on the fen's edge and the delicate beaked sedge (Rhynchospora capitellata) breezily waving in the center of the scene. Six months later, in March 2020, the pandemic had hit. As university classes pivoted online, we, as instructors, were forced to figure out how the unique shared experiences of the previous fall's classes, held in the field, could be translated meaningfully into a remote format for the upcoming summer and fall offerings. Field Botany and Taxonomy has been taught at the University of Rhode Island since the late 1940s. Professor Elmer Palmatier, a local botanical legend, established the class and was known to say: \"There should be no monotony when studying your botany.\" His legacy\u2014 students quickly learning hundreds of wild plants\u2014has been maintained by a lineage of memorable naturalists. Today, it continues in summer and fall classes led by Professor Brian Maynard, botanist Robin Baranowski, and their teaching assistants. The summer is an intense marathon to identify every plant found between late May and the end of June\u2014over 300 plants in a typical term. The sessions are composed of fast-paced, four-hour meetings, held four days a week. In the more traditional fourteen-week fall semester, the class heads out together twice a week to explore natural habitats around Rhode Island and identify about 250 plant taxa using sight, scent, texture, and even taste. Students collect and bind samples in herbarium presses for both courses and are constantly quizzed on plant names in the field. The courses cover both native and naturalized plants, with detailed units on grasses and mosses. The fall session becomes a race against time, given the threat of frost, and attention turns to autumn colors and winter twig characteristics. The coronavirus pandemic forced virtually all college courses online, many for the first time. Higher education as we knew it would change dramatically. While adequate tools for online education have been around for nearly two decades, most professors and students of the natural sciences had little experience with online learning, as it had never been necessary before. Now we had just a few weeks to move our courses entirely online before students returned from an extended spring break. Our most significant concern\u2014other than fears about keeping ourselves and our students safe from COVID-19\u2014was that we would not be able to provide our students with the quintessential field botany course experience. After much deliberation, we settled on a progressive learning structure that involved \"flipping\" the course. Instead of loading students up with plants to memorize through the usual sage-on-the-stage approach, we would hold the students responsible for finding and identifying plants on their own. While the traditional field course had emphasized learning a shared list of plants, this version would prioritize the development of skills that students could employ to identify any plant they encountered. Using an online learning platform called Brightspace, we created a series of modular lessons about the major groups of plants: wildflowers, trees and shrubs, ferns, and grasses. Each module included daily activities to train students on identifying the plants that they Field Botany in the Time of COVID-19 Emma Brown and Brian Maynard When Field Botany and Taxonomy at the University of Rhode Island went remote during the pandemic, the authors found that online tools like iNaturalist supported independent and flexible learning. This iNaturalist map from the summer term shows the wide distribution of class observations. MAP COURTESY INATURALIST; PLANT PHOTO BY SARAH MCDONOUGH found on their own. We centered these activities around multimedia tutorials on how to navigate four different field manuals (one for each major plant group) and two of the online keying systems found on the Native Plant Trust's GoBotany website. This was the first time we had used online keys for the class. The students would identify plants using the field manual or online keying system taught each week and then document their observations with photographs and notes using iNaturalist, a citizen-scientist app and website. These digital herbarium vouchers, as we call them, were formated according to a template we developed and took the place of the herbarium collection the students would have created for the inperson class. The new keying and vouchering skills of our students culminated in a capstone project. Each student designed a vegetation survey in a nearby natural area safely accessible during the pandemic. Students used iNaturalist to record the plants found along a transect line, pacing step-by-step and pausing at regular intervals to document the plant species encountered. The integration of iNaturalist into the class and requiring a vegetation survey were other firsts for the course. The summer session began in late May 2020 with eighteen students enrolled. Instruction was entirely asynchronous, meaning students could watch presentations and complete assignments on their own schedule. Students communicated with us by email, text, phone, and video calls. Challenges included making sure students had the necessary technology and access to natural spaces. We also needed to ensure that students understood the language of botany and, perhaps most importantly, that they could distinguish between native or naturalized plants and those in managed landscapes (which might not be found in their field guides). Fortunately, most students had smartphones that automatically tagged the photos uploaded to iNaturalist with GPS data. After keying and identifying a plant, the student would create a voucher with three clear images taken in the field and a description of the plant's shape, foliage characteristics, and other identification features. We guided students through the process of taking clear images. As a set, the photos should zoom to capture the entire plant silhouette, the branch arrangement, and finally up-close details of foliage, twigs, and flowers. Vouchers also included the steps used to identify the plant in the specified field guide, a link for that plant to the Consortium of Northeast Herbaria (a digital collection of herbarium sheets from dozens of herbaria), and an image of the plant on a plain white background with a digital herbarium label. The students posted the photos and notes to the class iNaturalist page, where the instructors, teaching assistants, and other iNaturalist users confirmed or challenged their identification. As new observations popped up on the iNaturalist map for the class, the difference from the in-person course was apparent. Instead of everyone learning the same plant in the same location, all in Rhode Island, we now racked up twenty-three unique records of sensitive fern (Onoclea sensibilis) from southern Maine to Philadelphia. One student reported plants sighted in a Maine salt marsh. Another documented vegetation in Manhattan parks. Each week, the students expanded one voucher into a presentation and posted it to a discussion blog. The presentations included a range map and notes on plant family characteristics, habitat, ecological relationships, and historical human uses. Blog conversations around these presentations became surprisingly animated: students enjoyed finding similarities in their plant-hunting adventures and learning new facts about plants they had also discovered, as well as about plants they had never seen before. Our learners went above and beyond our expectations by sharing photographs of the habitats and wildlife surrounding their botanical entries. Pictures of herons flew back and forth in the discussion posts, along with wild tales of adventurous plant-hunting escapades. Even a cinnamon-colored housecat participated in the fun as a model to show the size of cinnamon fern (Osmundastrum cinnamomeum) fronds against a large enough white surface for the digital herbarium voucher. These blog entries fostered engagement and interactions that we had thought were only possible in person, when we could walk back to the vans afer foraging cranberries, with fen water sloshing in our boots and conversations gushing. As it turned out, the blogs still allowed 4 Arnoldia 78\/4 \u2022 May 2021 the students to share their experiences with excitement and passion. In the last week of the class, the vegetation survey capstone tested the students' plant identification skills. After proposing a study area (which ranged from vacant lots to pristine forests), each student walked their transect and identified every plant species they found, posted their findings on iNaturalist, and produced a final report that they shared with the class. As the course unfolded, we found that the switch to the online format had created new learning opportunities. Students continued hands-on learning with greater independence. Resources designed for the course could be reused by students time and again, and we improved accessibility by captioning videos and narrating PowerPoints. Several students completed classwork from out of state, adding to the diversity of plants that the class found. The asynchronous schedule allowed students with personal or work obligations to participate fully. While our students all reflected that the course was time-intensive, they enjoyed the motivation to spend more time outdoors each week. After our success with eighteen summer students, we took stock of what worked best and ramped up for a fall course of fifty students. We ended up using many of the same tools developed for the summer class, but the material was now spread out over ten weeks and focused on the vegetation we would encounter in New England in late summer and fall. An added challenge of the pandemic was that students were scattered far and wide\u2014from Maine to Philadelphia\u2014 and could be forced into lock-down or quarantine at any time. For quarantined students, we prepared contingency samples, which included collections of photos and descriptions of habitat and plant characteristics that we observed in the field. While many fall students still attended remotely, we were finally permitted to meet in person, in small recitation groups, if students could get to campus. Twice a week, we helped up to five in-person students at a time with their keying and plant vouchers. We were initially concerned that students would learn only a fraction of the usual number of plants, but these concerns were assuaged by the depth of knowledge the students acquired Students created \"digitial herbarium vouchers\" for the class. Each voucher included at least three photographs of the plant in the field and one photograph showing the plant against a white background. ROBIN BARANOWSKI 6 Arnoldia 78\/4 \u2022 May 2021 for each plant and the confidence they gained in keying on their own. Across the summer and fall classes, our students posted nearly three thousand individual observations to iNaturalist\u2014about 360 unique species in each class. This number far surpassed the 300 or so plants taught in the past. Moreover, our students can now apply their plant identification skills anywhere in the world. We foresee that these tech-savvy citizen scientists will continue to use iNaturalist, including for BioBlitzes, which are intense twenty-four-hour events in which groups find and identify as many species of life as possible in a specific area. In explaining to our students how to learn their plants, we always stress that the best way to learn is to teach. The act of teaching others is a higher-level step in the learning process. The same students who initially had shied at the prospect of the online format shared plans to use their new knowledge for future careers and reported passing along what they had learned to friends and family. A select few students admitted to not liking plants before this class but noted that they learned to appreciate and even love the plants they encountered. Even as we return to in-person instruction this summer, we will use many of the tools we developed in 2020. We have committed to teaching a blended (online and in-person) field botany course to thirty-six students this fall. Moving forward, we expect to keep several of the teaching strategies that encourage independence and foster flexibility: keying modules, digital plant vouchers, a vegetation survey capstone experience, and the integration of iNaturalist and GoBotany. We are growing with the plants we teach. While the format may be different, the class is definitely a new sport off an old tree that we will continue to cultivate. For more information Visit our class iNaturalist sites at https:\/\/www.inaturalist. org\/projects\/uri-bio-323-summer-2020 and https:\/\/www. inaturalist.org\/projects\/uri-bio-323-fall-2020. GoBotany\u2014the Native Plant Trust's online tool for plant identification\u2014can be accessed at https:\/\/ gobotany.nativeplanttrust.org\/. This provided a valuable complement to the four field manuals that we also taught: Newcomb's Wildflower Guide by Lawrence Newcomb and Gordon Morrison, A Field Guide to Trees and Shrubs by George Petrides and Roger Tory Peterson, Northeast Ferns by Steve Chadde, and Grasses, Sedges, and Rushes by Lauren Brown and Ted Elliman. Acknowledgment Thanks to iNatauralist for permission to republish the map in this article. iNaturalist is a joint initiative of the California Academy of Sciences and the National Geographic Society. Emma Brown is completing her master of science degree at the University of Rhode Island and writing a thesis analyzing the experience of taking field-based courses online during the pandemic. This summer, she will return to her native Delaware, where she practices horticulture and compiles the Delaware Native Plant Society newsletter. Brian Maynard is a professor in the Department of Plant Sciences and Entomology at the University of Rhode Island. He teaches courses in plant propagation and production, landscape management, arboriculture, and field botany. Brian received the Gold Medal Award from the Massachusetts Horticulture Society in 2009 and the Award of Merit from the International Plant Propagator's Society in 2016. Students also submitted detailed notes with each digital herbaruim voucher. This section describes the steps taken to identify periwinkle (Vinca minor) using Newcomb's Wildflower Guide and also includes a link to a digitized specimen of this species at an herbarium. ROBIN BARANOWSKI"},{"has_event_date":0,"type":"arnoldia","title":"The Conference Must Go On","article_sequence":2,"start_page":7,"end_page":9,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25730","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25eb36c.jpg","volume":78,"issue_number":4,"year":2021,"series":null,"season":null,"authors":"Iles, Jeff","article_content":"The Conference Must Go On Jeff Iles Like a shimmering mirage on some lonely two-lane blacktop, the end of our global pandemic remained out of reach during the last academic year. No backyard barbeque with humans from another pod. No hockey games or theatre. No going anywhere sans facial covering. In my circle of fellow plant nerds, in-person trade shows and educational conferences topped the list of favorite social events that vanished. Remember those days? Striding up to the registration desk, receiving your official conference name badge, pawing through a complimentary tote bag filled with an eclectic assortment of swag, and then rushing off to the opening plenary session and, without giving it a second thought, sitting next to, or even shaking hands with, your randomly chosen seatmate. As 2020 dragged on and the 2021 conference season loomed on the horizon, it became abundantly clear to conference planners that inperson, traditional educational events were not a possibility, at least not for events scheduled for prime conference season between January and March. But the show must go on, right? This was my challenge as I contemplated strategies for keeping the flame alive for an educational conference I've managed since 1995: the annual Iowa State University Shade Tree Short Course, held on the university campus in Ames, Iowa. The event, which was heralding its sixty-fifth year in 2021, was the brainchild of Harold \"Sande\" McNabb, a forest pathologist at Iowa State. As the story goes, Dutch elm disease and its assault on our American elm (Ulmus americana) provided the impetus for the first gathering, which occurred at the McNabb residence. Now, many years later, the short course has become the can't-miss event for arborists and allied industry professionals in Iowa and surrounding states, drawing well over six hundred participants annually and featuring notable presenters like the late Alex Shigo, who encouraged us to \"touch trees\" and learn about their biology, care, and responses to wounding via compartmentalization. The themes, points of emphasis, and methods of instruction (handson workshops are always popular) vary from year to year. So, too, does the number of presenters (approximately thirty). But we never stray too far from discussing the benefits and maintenance requirements of these large, lifebreathing, woody friends. Not to overstate the importance of this conference or my hand in bringing it to fruition, but there can be no denying that the Shade Tree Short Course has earned its reputation as a trusted platform for arboricultural and horticultural education in Iowa and the upper Midwest. As the new year dawned, I felt an almost parental responsibility for the conference\u2014in part to continue McNabb's steadfast tradition, but also, even more importantly, to continue serving our loyal audience, some having attended since the late 1970s. Of course, our short course was not alone in facing this dilemma. Seemingly every educational conference around the country (even the world) was simultaneously confronted with the same set of circumstances and arrived at the same conclusion: \"If we're gonna do this, we're gonna have to go online.\" The world of video conferencing is a frightening place\u2014or at least it was for me. My fear was born out of the personal experience of witnessing even the simplest of virtual meetings with a handful of participants devolve into realtime lessons in frustration and futility. Who hasn't experienced the same? Poor or indecipherable audio. Low bandwidth prompting the meeting host to switch faces and voices into muted squares with names. Video conference platforms requiring tedious and sometimes confusing downloads\u2014and yet another password. If the downloads had required social security numbers and bank account information, I wouldn't have been surprised. Of course, I'm exaggerating for effect, but for those who grew up using technological advances such as the telephone, fax machine, electric typewriter, and those cute little personal computers (a.k.a., word processing machines) from the mid-1980s, 8 Arnoldia 78\/4 \u2022 May 2021 The Iowa State University Shade Tree Short Course is an annual conference that draws well over six hundred participants. In 2021, the event went online. JEFF ISLES Shade Tree Short Course 9 receiving a link that, if it worked, would transform desktop computers into portals to another realm could be a bridge too far. But what other choice did I have? Enter my grand plan. Historically, the Shade Tree Short Course takes place over two full days, but I knew that convincing an audience accustomed to working outdoors to stare at a computer screen for two solid days was going to be a nonstarter and, by extension, could have a dampening effect on attendance. Instead, I reasoned smaller chunks of virtual interaction and educational content would be far more palatable. Therefore, with wise counsel and advice from a university conference coordinator, we devised a week-long event at the end of February. Presentations would begin at eight in the morning and wrap up most days by eleven. Next, we needed to determine a fair registration fee for a virtual conference. Because I no longer had to worry about transporting and feeding my presenters, nor feeding participants, and because the number of educational sessions was reduced from previous years, I knew the registration fee used in 2020 ($170 early and $220 late) had to be reduced. With the intent of covering my remaining expenses (conference management fee and speaker honoraria), we decided on $40 for early registration and $55 for those coming late to the party. We also offered a reduced fee for university staff and students. But had I gone too far? In my attempt to provide an affordable product that would maintain registration numbers at least at a break-even point, had I committed the unforgivable sin of devaluing my own conference? As it turns out, full value for conference attendees was never in doubt thanks to the impressive lineup of speakers who, to a person, agreed at once to participate. And, to their credit, many graciously reduced or declined to accept their standard speaker fee, an acknowledgment perhaps of the reduced time commitment for a virtual conference. As the first day of the Shade Tree Short Course approached, however, one problem continued to silently orbit my conference, and its threat was potentially devastating: we needed to find the right video-conferencing platform. My unease was validated during a preconference practice session when our chosen video-conferencing platform performed in a less-than-satisfactory way. Most of my presenters were unfamiliar with the platform and found it user-unfriendly. When the same old audio problems surfaced, I knew it was time for plan B. Much to my relief, equipped with an alternate and reliable virtual conferencing platform and even a dose of unseasonably good late-winter weather (a nice touch even though we didn't need it), everything went swimmingly. No, we weren't able to offer the traditional scope of topics and workshops (over forty-five concurrent sessions spread over two days), but the aforementioned cadre of top-quality speakers made up for any deficiency in quantity. In the end, we attracted an audience of over 370 participants, including many longtime attendees and a few who'd never attended the short course before. In fact, many first-timers remarked that they attended in 2021 only because the program was offered online. And therein lies my next problem. Now that we've explored the realm of virtual education and witnessed its many benefits (the chat room was incredibly popular), many attendees would like our short course to preserve and integrate aspects of virtual programming in all future conferences. Ideally, a hybrid version could allow attendees to select from in-person sessions that would either be livestreamed or recorded for viewing later. In the end, cost and practicality will dictate the feasibility of such a hybrid model. Honestly, my preference would be for a return to our triedand- tested in-person roots; however, I also must allow for and accept that, in so many ways, the world has changed. This not-so-sudden immersion into the world of virtual conferencing has transformed the thinking of this reluctant conference chair. I now possess a new set of skills and have thoughtfully reconsidered what an educational conference should be. Just the same, while I can freely agree that learning doesn't necessarily require in-person, face-to-face interaction, virtual conferencing will always fall short as a replacement for engaging conversation around the coffee dispenser, in the buffet line, or gathered inside the pub at day's end. Jeff Iles is professor and chair of the Department of Horticulture at Iowa State University, in Ames, Iowa."},{"has_event_date":0,"type":"arnoldia","title":"Into the Valley of <i>Parrotia</i>","article_sequence":3,"start_page":10,"end_page":15,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25731","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25eb76f.jpg","volume":78,"issue_number":4,"year":2021,"series":null,"season":null,"authors":"Douglas, Phillip; Sj\u00f6man, Henrik","article_content":"The triumph and anguish of plant collectors can often be summed up with a single word: timing. No matter how well an expedition has been planned, collectors often confront either empty capsules or immature fruits. At other times, however, the fates align. In September of 2017, we embarked with colleagues on a collecting expedition to Azerbaijan, searching for multiple species poorly represented in botanical collections. The Persian ironwood (Parrotia persica) was our primary target, and for this species, our timing could hardly have been more auspicious. The Persian ironwood is an ornamental workhorse in the witch-hazel family (Hamamelidaceae) and is one of two species in its genus. Documented collections of Parrotia persica in public gardens tend to be from nurseries, and plants of known wild provenance are mostly sourced from populations in Iran. Although descriptions of the species' range tend to focus on the Alborz Mountains in northern Iran, plants do not typically recognize geopolitical boundaries, and thriving populations of Parrotia also exist in areas of the Hyrcanian forest and the Talysh Mountains of southern Azerbaijan. The flora in these biomes is considered a relict of a forest type that was much more widespread before glaciation events in the Quaternary, starting around two and a half million years ago. The Talysh region, in particular, includes more than ninety endemic species.1 Herbarium vouchers for Parrotia indicate a disjunct population in the country of Georgia, but it is widely believed these specimens were planted. In mid-September, our team departed the Azeri capital city of Baku and drove southward along the coast towards Lankaran. The trip had been organized by the Plant Collecting Collaborative, an organization consisting of eighteen botanical institutions, and our collaborators on the trip included Peter Zale from Longwood Gardens, Matt Lobdell from the Morton Arboretum, and Vince Marrocco from the Morris Arboretum. Vast agricultural fields dominate this landscape along the Caspian Sea, irrigated with the waters of the Kura River, which flows throughout the Caucasus region. Cotton, tea, grapes, and various citrus trees are the primary crops. Along the drive, we saw roadside plantings of Quercus castaneifolia, the chestnutleaved oak, which was another one of our species of interest. These plantings were the first we saw of the species in the country. After a long and bumpy drive, we were met in Lankaran by Hajiaga Safarov, deputy director of science at Hirkan National Park. Hajiaga committed his career to exploring southern Azerbaijan, documenting the flora and fauna. He graciously agreed to guide us over the next three days and assured us that he knew of several populations of Parrotia persica in the area. Departing from our hotel the following morning, Hajiaga led our team southwest of the city to the rural farming village of Az Filial. As we gained elevation, the paved highway soon ended, and we continued driving on a hard-packed, single-lane road. Cresting the top of a small hill, we suddenly found ourselves in the middle of Parrotia-dominant forest. Scant herbaceous vegetation existed under the canopy of these magnificent trees, a result of intense grazing pressure from the surrounding farms. We parked under the shaded canopy of ironwoods and began to hear tapping on the car's roof, as though a light rain were passing over. The cloudless sky was not precipitating; the sound we heard was something much more miraculous. Plants in the witch-hazel family exhibit a unique form of seed dispersal. As the capsules of Parrotia persica begin to dry, the exterior walls (technically the exocarp) shrink in size and begin to apply pressure to the seed, causing its forceful ejection. This method of seed dispersal\u2014the so-called drying squeeze catapult2\u2014was the source of the light raining sound. When we exited our vehicle, we witnessed small, black seeds bouncing off the roof Into the Valley of Parrotia Phillip Douglas and Henrik Sj\u00f6man DOUGLAS, P. AND SJ\u00d6MAN, H. 2021. INTO THE VALLEY OF PARROTIA. ARNOLDIA, 78(4): 10-15 The Persian ironwood (Parrotia persica) fills a valley near Lerik, Azerbaijan. When the authors first encountered this overlook in 2017, the diversity of fall color and form was unmistakable. This photo was taken on a return trip in 2019. ALL PHOTOS BY PHILLIP DOUGLAS UNLESS NOTED and hood. In a marvelous turn of fate, we had timed our trip to document and collect Parrotia at the most advantageous time. Witnessing the forceful ejection of these seeds only added to the intrigue of the species. All hands worked quickly to obtain fruits that had not yet dehisced. We gathered several hundred capsules from throughout the population. Diversity in the Wild The Hyrcanian forests extend from southern Azerbaijan into Iran, wrapping around the southern coast of the Caspian Sea. In Azerbaijan, Parrotia occurs at elevations between sea level and around 1,600 feet (500 meters). Strong cultural influences of forest grazing, active felling of trees for firewood, and coppicing for fencing materials and winter feed have transformed the landscape. Farmers also coppice trees to minimize the shading of valuable meadow environments that provide winter fodder for sheep, cattle, and goats. The extensive coppicing in this region has made it difficult to see the natural habitat and variability of Parrotia. Examining the approximately fifty trees within the small population that we first encountered, it quickly became clear that an impressive amount of genetic variability was present. Bark characteristics alone were distinctly different, with variation including creamy, dappled camouflage mottling and golden, iridescent, paperthin flakes. It was far too early in autumn to see any fall color in this population, but we suspected that variation might exist for this trait as well. After making another collection from a heavily fruited Caucasian zelkova (Zelkova carpinifolia), we departed from the site and headed farther south towards the Hirkan National Park. Driving along the Lerik-Lankaran highway, we saw the Talysh Mountains begin to slowly build elevation as the forested areas became more dispersed between meadows and xeric terrain. Hajiaga was leading us to a historic cemetery and mosque outside the village of Babagil. In addition to Parrotia, our group was targeting several other unique woody species: the chestnut-leaved oak and a subspecies of the common boxwood that is endemic to southern Azerbaijan, Buxus sempervirens subsp. hyrcana. We encountered both species outside of the cemetery and mosque. This site dates to the sixteenth century and contains many enormous planted specimens of Caucasian zelkova and chestnut-leaved oak. Across the road from the cemetery is a remnant piece of the The first population of Parrotia persica that the authors visited in Azerbaijan revealed a typical, overgrazed understory. Yet the trees displayed variable and unique bark. Parrotia 13 Hyrcanian forest. Here, we discovered large boxwood growing in the heavy shade of Parrotia persica. Just beyond the roadway, we encountered our first large specimens of the chestnut-leaved oak. They created a towering forest canopy over 65 feet (20 meters) tall, with trunk diameters reaching over 3 feet (1 meter). Unfortunately, these two species develop seed at the opposite ends of autumn; the boxwood had already dehisced, and the oaks were not yet ripe enough for collection. We were able to make a large collection of intact seed capsules from the Parrotia on the property. This collection, at 1,510 feet (460 meters), marked the highest elevation at which we found Parrotia growing, and it should make for an interesting evaluation for cold hardiness. Departing westward, our group continued towards Lerik, a historic mountain town perched at 3,600 feet (1100 meters), overlooking the border with Iran. Gazing southward from the windows of our vehicles, we came across a magnificent sight: a sprawling forest of Parrotia persica filled the expansive valley beneath us. Towering velvet maple (Acer velutinum) dominated the upland areas, and enormous Caucasian alder (Alnus subcordata) were dotted along a slow-moving creek. Azerbaijan had been plagued in 2017 with a major drought, leaving the herbaceous layer completely dormant in autumn and adversely affecting the quality of autumn color. Despite this drought, the Parrotia in this valley showed deep hues of burgundy, red, orange, and yellow. Throughout this population, a diversity of form was also present. We noted many trees with dense conical crowns and a strong branching hierarchy. These structural characteristics would be well suited for trees selected for urban plantings. We were unable to access the forest because we had much more work ahead of us, but the memory of this valley remained with us after the trip. A Return to the Valley In late October 2019, the two of us traveled again to Azerbaijan to attempt collecting the chestnut-leaved oak from throughout its northern range. Similar to Parrotia persica, this species only occurs in the mountains of southern Azerbaijan and northern Iran. Its acorns don't fully ripen until late in the season, and we hoped to collect them before they fell to the ground, where insects and herbivores can render them useless. The drive south from Baku to Lankaran took half of the time during this trip, as construction of a multilane freeway had been completed, connecting Baku to Tehran, Iran. Our failure to collect acorns from this rare oak had haunted us for the past two years, and we were eager to determine if we had properly timed our trip. The landscape throughout southern Azerbaijan looked vastly different compared to 2017. Precipitation had fallen evenly through the year, and the previously dormant herbaceous layer was putting on an amazing show. The meadows surrounding the Babagil cemetery and mosque were filled with flowering geophytes. Two species of crocus (Crocus speciosus and C. caspius) carpeted the landscape and appeared almost as a monoculture lawn in areas that were heavily grazed. Pink-flowered cyclamen (Cyclamen coum) dotted the shaded understory of the endemic boxwood. The flowering spectacle was a wonderful sign of good seed development, and we were able to make three separate collections of chestnut-leaved oak at elevations ranging from 1,540 to 2,900 feet (470 to 900 meters). After finishing our oak collecting early, we had time to indulge in the forests of Parrotia persica. As we drove along the highway from Lerik, back to our accommodations outside of Lankaran, we made a familiar stop to gaze across the valley of Parrotia that we had discovered two years before. Our timing was once again rewarded with amazing views of the valley in full autumn colors. It is difficult to describe the array of colors. Individual trees within the canopy exhibited shades of deep burgundy, brick red, orange, and buttery yellow. We decided to use our remaining day of the trip to attempt to access and document this population. We collected GPS coordinates and headed back to our accommodations to plan the next day's work. After looking over various maps and satellite images, we were able to devise a way to drive as close as possible to the ridgeline across the valley, where several small houses stood. Our goal was to closely examine the trees in this population, taking photographs to document autumn color and differences in form. Trees The authors ventured into the valley of Parrotia in late October 2019. Fall color took on rich variation. Trees with dense, pyramidal habits (left) suggest exceptional potential for urban plantings. Phillip Douglas (bottom right) stands with a large Parrotia observed at another location earlier in the trip. HENRIK SJ\u00d6MAN Parrotia 15 with exceptional qualities would be geotagged so that we could return to them for propagation material in the coming years. The following morning, we departed the hotel and headed towards the valley, excited by the prospect of getting to walk beneath the canopy of the relict forest. The paved road quickly turned into a dirt path, and after crossing over a shallow creek, it became a deeply rutted, muddy quagmire. Our translator and driver, Ilgar Guliyev, guided us through the terrain with expert precision. We soon found ourselves parked outside of a small farmhouse, and Ilgar went in to inquire about accessing the valley below the property. After a short conversation with the owners, we were informed that the valley belonged to the state, and our collecting permits would allow us access to the site. Basing our navigation on several massive chestnut-leaved oaks and oriental beech (Fagus orientalis) along the top of the ridge and a group of towering Caucasian alder at the bottom, we began traversing towards several Parrotia we had photographed the day before. The first selection that we documented exhibited a uniform, brick-red autumn color throughout the canopy. We continued to traverse up and down the steep slopes of the hill, documenting selections with peachy-pink autumn color, dense and pyramidal habits, and even dappled burgundy and green foliage. The diversity of the species within this singular valley was amazing to see. We hope to return to the valley in late spring to obtain scion wood from these selections to begin growing and evaluating their performance in various climates and conditions. From the Wild, Into Cultivation The study and documentation of plants in situ is a valuable means of determining species that are well suited for urban horticulture and other specific uses. In Lankaran, we were also able to see how Parrotia persica has been used locally in extensive urban plantings. The species could be seen in park environments as well as in small curbside planter spaces. The hot, dry summers of Lankaran coupled with challenging site conditions of urban environments did not seem to affect this highly adaptable species. As a street tree, the species often becomes too wide, resulting in unflattering pruning efforts, but this issue could be solved with more intentional selection. As we had observed, an extensive variation in the size and expression of Parrotia occurs in the wild, suggesting the fantastic development potential of the species for public plantations in both Europe and North America. In cultivation, Parrotia is mainly represented by seed-propagated material, which results in large variations, making it difficult to predict mature size and habit. Presently, cultivars of Parrotia persica available on the market include 'Vanessa', 'Ruby Vase', and 'Persian Spire', which all represent narrow-growing forms. Based on our field observations, the species has significantly more expressions that deserve to be evaluated in cultivation. We hope to develop new cultivars of this species that will have uniform size and fall color characteristics. The species' adaptability to periods of intense heat and dry soil conditions, coupled with its tolerance for high pH soils, makes it a perfect candidate for further development as an urban tree. Hopefully, we will once again be blessed with perfect timing to collect from these populations and continue working with this relict species. Endnotes 1 Safarov, H. M. 2009. Rare and endangered plant species in Hirkan National Park and its environs. In N. Zazanashvili and D. Mallon (Eds.). Status and protection of globally threatened species in the Caucasus (pp. 193-198). Tbilisi: CEPF, WWF. 2 Poppinga, S., B\u00f6se, A. S., Seidel, R., Hesse, L., Leupold, J., Caliaro, S., and Speck, T. 2019. A seed flying like a bullet: Ballistic seed dispersal in Chinese witch-hazel (Hamamelis mollis Oliv., Hamamelidaceae). Journal of the Royal Society Interface, 16(157): 1-10. http:\/\/ doi.org\/10.1098\/rsif.2019.0327 Acknowledgment This work would not have been possible without the guidance and expertise of our partners at the Azerbaijan National Academy of Sciences, Institute of Botany and Central Botanical Garden of Azerbaijan, and the Hirkan National Park of the Ministry of Ecology and Natural Resources, Republic of Azerbaijan. Funding was generously provided by the Daniel F. and Ada L. Rice Foundation. Our work is dedicated in memoriam of Dr. Hajiaga Safarov (March 1, 1963-November 17, 2018). Phillip Douglas is the director of plant collections for the Chicago Botanic Garden and also serves as the chair of the Plant Collecting Collaborative. Henrik Sj\u00f6man is a senior researcher at the Swedish University of Agricultural Sciences and the scientific curator at the Gothenburg Botanical Garden."},{"has_event_date":0,"type":"arnoldia","title":"","article_sequence":4,"start_page":16,"end_page":23,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25732","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25ebb27.jpg","volume":78,"issue_number":4,"year":2021,"series":null,"season":null,"authors":"Elkin, Rosetta S.","article_content":"Barrier islands are young landscapes. Although absolute dates are hard to pin down, the barrier islands that ring Florida's coast are only about five thousand years old and represent some of the most dynamic landscapes in the world. In the context of earthly timescales, the islands surfaced at the end of the Stone Age, around the same time that written language was developed in Ancient China and humans began to interact with yeast microorganisms for producing alcohol and bread. At the time, plant life was already well established for millions of years, taking root firmly and resolutely across landscapes that were only slightly more intact than not. Today, Florida's coastline extends 1,350 miles, of which 700 miles are structured by barrier islands that are characterized by urbanization rather than earthly formation. Development is intended to prevent the young landscape from further formation, arresting worth in property value while securing costly infrastructure projects. Young soils are paved and only tend to host disturbance-adapted plants that creep in along built lines, chain-link fences, beachfront terraces, and in the obvious cracks between sidewalks. The most iconic plants are the mangrove species (Rhizophora mangle, Laguncularia racemosa, Avicennia germinans) that silhouette the shoreline, while florific beach sunflowers (Helianthus debilis), green-fruited pond apples (Annona glabra), and sea grapes (Coccoloba uvifera) with dense crowns are commonly found inland. In this setting, few remnants of the barrier island ecology remain amidst the rich imported flora of the mixed tropical and temperate zones. If you consult a map of Florida on your handheld device, the string of thin barrier islands that contour the coast is barely legible. Zooming in yields more clarity between land and water. Each barrier island floats along the shore of the mainland, stitched together by a line of causeways and interstate roads that seem to pull the islands landward, or stop them from moving seaward. Now, zoom in on the west coast near Fort Myers. Here, the stitch is called the Sanibel Causeway, which starts at a small crossing known as Punta Rassa. The causeway is supported by a sandy spit that separates Pine Island Sound from the Gulf of Mexico. The route extends into Periwinkle Way and stretches the length of Sanibel until it turns into the next stitch line at Blind Pass, a managed inlet known for shelling and fishing. Blind Pass is the last stop before arriving on Captiva Island. Consider the same map, and zoom in again on Captiva Island: the gray asphalt of parking lots and sidewalks, the vectorized streets and alleys, and the blank fills of the private space around each foundation. If you search for directions, the route leads you past green golf courses and beige beaches, while the rest of the landscape is defined by different shades of gray. There is no public information beyond the built form, and certainly no recognition of plant life. The lack of public knowledge about plants always strikes me as unusual, although it comes up frequently in my work as a practicing landscape architect and as a professor and An Impermanent Inventory: Plant Collections for a Changing Climate Rosetta S. Elkin \"Permanence doesn't really interest me. My whole focus has been on the activity of my life. Out of the activity has come a mass of works, which are really just evidence that I'm still paying attention.\" \u2014Robert Rauschenberg Facing page: Captiva Island, on the southwestern coast of Florida, is especially vulnerable to the effects of climate change, including sea-level rise. In 2017, the author was commissioned to develop a landscape-adaptation plan for the former home of Robert Rauschenberg on Captiva. A dynamic plant inventory would be essential. ALL IMAGES COURTESY OF THE AUTHOR ELKIN, R. S. 2021. AN IMPERMANENT INVENTORY: PLANT COLLECTIONS FOR A CHANGING CLIMATE. ARNOLDIA, 78(4): 16-23 18 Arnoldia 78\/4 \u2022 May 2021 researcher, studying the interactions between human and plant life. Within landscape architecture, the prominence of pathways and built structures seems to resonate with the public more than careful attention to particular plants. Presumably, this is one reason why landscape architecture is losing plant knowledge.1 So when it comes to finding your way in a new landscape, it is no wonder that the only means of tracking distance and not getting lost are found in the gray surfaces that demarcate outward appearance and built materials. But, as streets are inundated, seawalls fail, and foundations erode, might the endurance of plant life be appreciated in new ways? Designing a Plant Inventory In 2017, I was commissioned to study the changing conditions at the home of Robert Rauschenberg on Captiva Island, in order to propose a landscape-based adaptation plan to the effects of a changing climate.2 These effects include, but are not limited to, sea-level rise. Across Florida, the effects cascade: warmer waters increase the velocity of hurricanes, increased salination threatens drinking water supplies, the blooms of red tide devastate sea life, while blue-green algae amalgamate with heavy erosion to suppress tourism. The risks brought on by our warmer climate are not singular, which is why there is no simple solution. Rauschenberg cared deeply for Captiva both in terms of creative inspiration and also because it appealed to his ideas of impermanence, so elegantly stated in an interview about his art process: \"Permanence doesn't really interest me.\" When we were guided through our first site visit, intricacies of the built landscape were prioritized, including workshops for printmaking and dance studios, a beach house, the main studio, and the historic Fish House\u2014a building perched in the bay.3 Yet, the grounds are most remarkable because they encompass twenty acres of uninterrupted barrier island, a landscape that bridges the bay and the beach sides. Most properties either enjoy views of the beach or the bay, but rarely both. The Rauschenberg campus is verdant and alive with a continuous canopy that distinguishes it from the rest of the island because Rauschenberg valued the dynamic landscape and never sought to arrest and define it. The grounds\u2014now used to host an internationally recognized artist residency program\u2014are so culturally rich and ecologically lively that there was no lack of inspiration, and I was eager to get started. At its widest, Captiva is two thousand feet wide; at its narrowest, only about four hundred feet. The Rauschenberg campus sits along the widest portion. Despite its verdant ecology, a standard map registers gray tones, presumably because private land is not rendered beyond building footprints. As the project began, I sought more detail from standard site plans and surveys, the basis of architectural traditions, anticipating more specificity because Rauschenberg himself was so committed to his plants. In particular, he was committed to maintaining an area that he called the jungle, a ramble of sprouting spontaneous plants that makes up almost half the site.4 Rather, we were handed a site plan that outlined the property lines and included the building footprints, connected by a path system. The rest of the site was white. A site plan without any indication of plants is not only blank; it creates the impression of a landscape devoid of life. As a result, our first act of design was to put the plants back on the map. Creating a plant inventory for a landscape architectural project is not a normative or established convention. But a plant inventory is a curatorial tradition that supports research within the living collections of arboreta and botanic gardens. An inventory charts long-term change and unlocks the puzzles of horticulture, so it is surprising that inventories are not more of a standard in professional practice. The objective of a plant inventory is to document and describe the current status of a collection. Over time, the inventory can be compared to past iterations, revealing landscape changes.5 In turn, this secures a plan for future plantings. A plant inventory must be updated in order to remain dynamic, which requires ongoing interaction in the field. This is especially true because plants move, die back, transform, and sometimes shift from their original locations. Captiva Island Inventory 19 Typically, an inventory is established at the same time as a garden and creates a baseline to determine future accessions and deaccessions. For instance, the first accession records at the Arnold Arboretum date to 1872, the year the institution was founded, although it took about a decade for the initial card-file system to be refined. In an account from 1881, Charles Sprague Sargent outlines the importance of the inventory but admits that accurate records are often abandoned because they are \"too expensive for practical working.\"6 He references the future value of recording each plant despite the challenge, suggesting that the effort must bear the test of time. At the Rauschenberg campus, our team believed that the strain of changing climates made the connection to time even more powerful. Establishing a curatorial tradition within an undocumented collection posed two important challenges to the inventory from the start: first, to establish what constituted a \"tree\" among a host of woody plants, and second, to assess a largely spontaneous collection. Both challenges forced us to make value judgments based on what to count, and thus what to omit, a puzzle that raised more questions than we could answer alone. The Inventory Process The Sanibel-Captiva Conservation Foundation (SCCF) was founded in 1974 by a group of Islanders committed to the preservation of the island ecosystem. At the time, SCCF successfully opposed development in Sanibel by incorporating as a city, enabling votes on dredge-and-fill policy, uprooted mangroves, seawall construction, and overscaled condominiums. 7 The same constituency hired the firm of Ian McHarg, the renowned landscape architect who wrote Design with Nature, an Initial site plans and surveys for the Rauschenberg campus emphasized the built infrastructure. Notably, the plants were unrepresented, even in the densely vegetated area known as the jungle. Captiva Island Inventory 21 influential ecological treatise.8 Captiva did not follow suit and has experienced the consequences of haphazard planning ever since. This is one of the main reasons that the Rauschenberg campus is so uniquely important: it is an anomaly in the landscape that might help inform Captiva's future. Our team, based in Massachusetts, worked with local horticulturist Jenny Evans from SCCF to initiate the process of developing a baseline for the plant inventory. Without a baseline, neither preservation nor conservation exists. It creates a reference for measuring and assessing disturbance. Although Jenny and her team had little experience establishing a plant inventory, she saw value in the challenge due to the extremities of change expressed by plant loss throughout the hurricane season. The baseline would help us chart the rapidity of change in both the loss of material in hurricane season and, hopefully, the regrowth of disturbance-adapted species. Collectively, we were motivated to tackle the questions raised about the process of gathering and digitizing the data because we saw the importance of creating publicly accessible plant knowledge. Our inventory would prioritize woody plants, but as we worked through our initial questions, we found that trying to define a \"tree\" at Captiva proved conceptually hazardous in itself.9 Many woody plants do not behave as trees with a single trunk, but clump or spread. To capture this distinction, we created two categories of data: rather than discriminating between trees and shrubs, we suggested points and areas. Points recorded the center of woody plants with single trunks. Areas recorded the total diameter of the woody plant\u2014the perimeter of all trunks and shoots. Each point was recorded in a discrete location using latitude and longitude, while areas were recorded by walking the perimeter of the plant and recording the path.10 The system of areas was especially useful for taking stock of the mangrove fringe on the bay side, yet flexible enough to allow us to indicate where specific points were noticeable as major trunks within the tangle. The points within the mangrove area are only one example of how the standards of defining a tree helped us standardize a method across a site full of exceptions. As trees were defined and included in the inventory, a workflow developed between the on-site project team and the data input team. First, the site was divided into 75-by-75-foot quadrants in order to work systematically across the landscape. The quadrants did not have to be delineated in physical space: they were charted by datasets of a handheld GPS device. The on-site team then recorded woody plants using the system of points and areas, and the data from each quadrant was shared with our team sitting at our studio in Massachusetts. This workflow enabled the field team to move from one quadrant to the next and continue to amass data.11 Our team uploaded their new field data to a global information system (GIS) and aligned this work with site surveys used in the original design documents.12 We checked the data, cleaned duplicates or errors, and assigned a unique catalog code in GIS, which was exported with labels and integrated into the site survey. The process raised questions about what type of data was most useful to contain on the map label and how the information could be read by those both familiar with and unfamiliar with plants. Therefore, we decided on two distinct categories: standard and custom. Standard data included common, Latin, and family names, along with trunk diameter (at breast height) in centimeters, height taken in meters, geospatial location (latitude, longitude), location on site (quadrant), and the year recorded. To include canopy cover in the standard category, Jenny came up with a novel expression\u2014a range from one to five\u2014that corresponded to how much of the sky could be seen when standing at the trunk. If 80 to 100 percent of the sky was obscured, she would give the canopy a five; 60 to 80 percent obscured would be a four, and so on. This might not seem relevant in the context of temperate trees, but in a tropical site that is largely overgrown by densely sprouting palms, the canopy can still lack density, which affects overall shade and comfort despite height and maturity. We also assigned a Florida Exotic Pest To develop the plant inventory at the Rauschenberg campus, a field team collected GPS points, measurements, and detailed observations for all woody plants growing on the twenty-acre property. The complete inventory can now be accessed on a handheld device. 22 Arnoldia 78\/4 \u2022 May 2021 Plant Council category to each plant. Finally, we created a unique identifying code for each woody plant in the inventory. The custom category necessitated the most creative collaboration as we imagined what future residents and stewards might wish to know about the plants of the present. The first section within the custom category includes descriptions of environmental influences (damaged or broken limbs, leaning habit, and so forth), notes about neighboring plants in relation to the spread (consider for instance Ficus aurea, the strangler fig, which envelops a host tree), and surveyor comments. The collaboration with SCCF was crucial to the comments section and includes remarks about character or significance that were personal, such as \"never seen it grow this way\" or \"covered in lianas,\" a crucial input to research in heavily urbanized landscapes that resist standards. The subsection also provides space for more nuanced assessments of the Florida Exotic Pest Plant Council criteria, with notes such as \"typically invasive, but not aggressive on this property\" that overcome the binaries of what typically counts and what doesn't count in a living collection. In the Context of Change Landscape design often implies stability and predictability. Yet, the dynamics of the landscape are changing, which invites practices to change in turn. This need is especially pronounced on the Florida coast. As we looked for models for our project, we consulted with curatorial staff at public gardens and found a range of concerns. At the Arnold Arboretum, for instance, staff pay especially close attention to evidence of infestations, as some of the most devastating losses to the living collection are brought on by foreign pathogens.13 While the rise of foreign pathogens is certainly not bound to the Northeast, Florida must first contend with the intensely localized effects of increased storm damage brought on by rising seas. A more apt comparison might be made to the inventory at Montgomery Botanical Center in Coral Gables, Florida, a historic collection specialized in the conservation of palms, cycads, and conifers from across the world. The garden is a coastal site vulnerable to episodes of increased storms and the very real effects of about one-third of an inch (nine millimeters) of rise in sea level per year.14 Thus, Montgomery is grappling with a concern common to all coastal living collections in a time of rapid climate change: How far into the future should we plan? While this is an enduring question in relation to living collections, it finds amplified resonance considering that Montgomery calculates an increased inundation of forty-three acres, or 36 percent of the entire garden.15 While this number is staggering, the plant inventory confirms that only 8 percent of the collection will be lost in this scenario. Although the figure does not include storm damage, salt intrusion, and other vulnerabilities, it does significantly change the answer to the question: planning can no longer occur in one-hundred-year increments. The status of any living collection is dependent on maintaining an inventory, which raises questions as to why plant inventories are not more commonly practiced beyond the world of public gardens. In the context of barrier islands, like Captiva, change is noticeable seasonally as hurricanes sweep across the surface of the land while fluctuating sea levels remake the coastline. But, of course, landscapes everywhere are increasingly in states of flux. The knowledge of how to create and maintain an inventory is critical to engendering a unique collaboration between plant and human life within our everyday landscapes. A plant inventory is a record of human and biotic adaptation, a neutral middle ground that accumulates experience and data. It helps visually connect the public to the effects of accelerated climate change, and in a practical sense, it inspires care and helps humans take notice of the plants in their environment. After the success of developing the plant inventory at the Rauschenberg campus, our team's ensuing idea is to adapt the same open-source technology into a handheld, userfriendly platform that could form the basis of a public inventory for landscapes anywhere, populating our blank site plans and challenging generic street views. We imagine citizen scientists learning to create a site history, as plants under their stewardship become a baseline for future generations. Plant inventories are cruCaptiva Island Inventory 23 cial to increasing an awareness of change, especially in the face of both chronic and episodic stresses of the twenty-first century. Perhaps we can shape an understanding of change by visualizing and valuing impermanence. Endnotes 1 A number of authors, myself included, write about the loss of plant knowledge in design. See, for instance: Raxworthy, J. and Harrisson, F. 2018. Overgrown. Cambridge: MIT Press. 2 Practice Landscape includes Emily Hicks and Joanna Lombard, and we were commissioned by the Robert Rauschenberg Foundation to work as part of a team in collaboration with WXY architects and eDD engineers. 3 Rauschenberg bought the Fish House from Jay Norwood \"Ding\" Darling, chief of the US Fish and Wildlife Service (formerly the Biological Survey). Ding Darling is best known for ushering in the Federal Duck Stamp Program to expand the federal purchase of wildlife habitat. See, for instance: Ding Darling Wildlife Society. n.d. Our namesake. https:\/\/ dingdarlingsociety.org\/articles\/our-namesake 4 The cultural history of the plantings is culled from various oral accounts and conversations, especially with Matt Hall, the site manager who worked closely with Rauschenberg on Captiva, until Rauschenberg's passing in 2008. 5 The Arnold Arboretum plant inventory claims that to meet objectives \"the Arboretum fields expert curatorial staff able to conduct inventories as well as troubleshoot an array of taxonomic, cartographic, and horticultural puzzles.\" See: Arnold Arboretum of Harvard University. 2011. Plant inventory operations manual (2nd ed.). http:\/\/arboretum.harvard.edu\/ wp-content\/uploads\/2020\/07\/plant_inventory_ operations_manual.pdf 6 Sargent, C. S. 1882. In Harvard University, Annual reports of the president and treasurer of Harvard College, 1881-82 (pp. 122-123). Cambridge, MA: University Press. 7 SCCF's mandate continues to advocate through education and outreach, supported by an intellectual generosity and a spirit of collaboration. For a short history of SCCF in the context of early development see: Davis, J. E. 2017. The Gulf: The making of an American sea (pp. 406-410). New York: Liveright Publishing Corporation. 8 McHarg, I. L. 1969. Design with nature. Garden City, NY: Published for the American Museum of Natural History [by] the Natural History Press. 9 We initially turned to a definition of trees provided by the Arnold Arboretum's Peter Del Tredici: \"A tree can be defined as a plant that, when undisturbed, develops a single, erect woody trunk. A shrub, on the other hand, is a woody plant that, when undisturbed, branches spontaneously at or below ground level to produce multiple stems. In general, a tree will develop secondary trunks in response to injury to its primary trunk or root system, to displacement of its primary stem out of the normal vertical orientation, or to a dramatic change in surrounding environmental conditions.\" Despite the usefulness of this definition, in practice, we found the distinction was difficult to apply at Captiva. Del Tredici, P. 2001. Sprouting in temperate trees: A morphological and ecological review. The Botanical View 67: 121-140. 10 Data was collected using a handheld Trimble, a GNSSbased data collector that is integrated with ArcMap GIS and is the standard in forestry surveys. This system allows for ease of data entry and storage that works well with our needs for both quantitative and qualitative data. Model: Trimble Geo 7X. 11 The field team received the initial GIS data for each quadrant as a CSV and shapefile. 12 This data alignment involves changing the coordinate system to a projected coordinate system. 13 Emerald ash borer (Agrilus planipennis) and hemlock wooly adelgid (Adelges tsugae) are of particular concern in eastern Massachusetts. Among numerous scientific studies on monitoring, see, for instance: Knight, K. S., Flash, B. P., Kappler, R. H., Throckmorton, J. A., Grafton, B., and Flower, C. E. 2014. Monitoring ash (Fraxinus spp.) decline and emerald ash borer (Agrilus planipennis) symptoms in infested areas. General Technical Report NRS-139. Newtown Square, PA: United States Department of Agriculture, Forest Service, Northern Research Station. 14 Wdowinski, S., Bray, R., Kirtman, B. P., and Wu, Z. 2016. Increasing flooding hazard in coastal communities due to rising sea level: Case study of Miami Beach, Florida. Ocean & Coastal Management, 126: 1-8. 15 According to a one-hundred-year projection: Griffith, M. P., Barber, G., Tucker Lima, J., Barros, M., Calonje, C., Noblick, L. R., Calonje, M., Magellan, T., Dosmann, M., Thibault, T., and Gerlowski, N. Plant collection \"half-life:\" Can botanic gardens weather the climate? Curator: The Museum Journal, 60(4): 395-410. Rosetta S. Elkin is an associate professor at McGill University, an associate of the Arnold Arboretum at Harvard University, and the founder and principal of Practice Landscape. Rosetta's work considers living environments with a particular focus on plant life and climate change. She teaches planting design, fieldwork, and seminars that advance a theory of plant life between ecology and horticulture."},{"has_event_date":0,"type":"arnoldia","title":"William Purdom: The Forgotten Arnold Plant Hunter","article_sequence":5,"start_page":24,"end_page":37,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25733","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25ebb6b.jpg","volume":78,"issue_number":4,"year":2021,"series":null,"season":null,"authors":"Gordon, Francois","article_content":"William Purdom spent three years collecting in northern China and Tibet on behalf of the Arnold Arboretum and the British nursery James Veitch & Sons. Here, Purdom passes through a gate in the Great Wall, in Shanxi Province, in the spring of 1910. PHOTO: \u00a9 BRITISH LIBRARY BOARD It was early March 1912, on the banks of the Yellow River, 450 miles south of Beijing. An Arnold Arboretum plant collector and his three-man escort had ridden more than five hundred miles east from Minxian, in Gansu Province, through a region devastated by the Xinhai Revolution. The revolution had toppled the last Qing emperor and replaced the centuries- old imperial system of government with a republic, which was struggling to establish its authority against a plethora of regional warlords. The roads were alive with bandits, and food and shelter hard to find, but the collector's journey to date had been uneventful. He and his escort were drawing near their destination, the railhead to Beijing in Honan (now Luoyang), the provincial capital of Henan Province. Suddenly, they were ambushed by a group of mounted men, who fired as they charged, killing two horses in the first moments of the attack. It's unlikely that the bandits knew what the travelers' saddlebags and packhorses' loads comprised, still less that they coveted the herbarium specimens and the seeds and tubers laboriously collected in Gansu and Tibet over the previous year. But a foreigner was sure to be carrying silver specie to pay his way on the road, and the surviving horses would fetch a good price. The botanist, however, had other ideas. He drew a lever-action rifle from the scabbard beside his saddle and, as he would later write, \"made a stand,\" shooting three of the attackers and several of their horses. His escort joined in, driving off the bandits, and the party galloped to the small city of Shenchow, from where they eventually continued their journey to Beijing, which passed without further incident.1 The plant collector was William Purdom, at the conclusion of a three-year expedition on behalf of the Arnold Arboretum and the British firm of James Veitch & Sons to northern and northwestern China and the Tibetan region of Amdo. In the course of his expedition, he sent to Boston 550 packages of seeds and well over one thousand herbarium specimens.2 Purdom, born in 1880, was a head gardener's son from the Lake District in northern England. He served an apprenticeship with his father before working for two distinguished London nurseries, then joining the Royal Botanic Gardens, Kew, as a student gardener. The Kew course of training for botanists and horticulturalists was internationally renowned and correspondingly demanding to join and to pursue. Purdom had done well and had proved a particularly skilled propagator, especially of woody plants. But Kew's director, Sir William Thiselton-Dyer, did not appreciate Purdom's activism as the secretary of the Kew Employees Union, and in 1905, Purdom was dismissed for \"agitation.\" Purdom promptly petitioned the Board of Agriculture, Kew's parent ministry, which agreed that he was perfectly entitled to join a trade union and ordered his immediate reinstatement. Thiselton-Dyer, unable to bear this humiliating public reversal, resigned. The new director, Colonel David Prain, then had to contend with the only strike there has ever been at Kew, efficiently organized by Purdom. All in all, it's perhaps not surprising that when, in 1908, Charles Sprague Sargent enquired whether Kew could recommend someone to undertake a three-year expedition to China, Prain enthusiastically recommended Purdom as the very man for the job!3 Sargent had come to Britain in August 1908 to engage a plant collector to travel to northwestern China to collect plants and seeds for the Arnold Arboretum. Ernest Wilson, whom Sargent had sent to China in 1907, had made it clear that he would not extend his two-year contract.4 In 1906, Sargent had also agreed William Purdom: The Forgotten Arnold Plant Hunter Francois Gordon GORDON, F. 2021. WILLIAM PURDOM: THE FORGOTTEN ARNOLD PLANT HUNTER. ARNOLDIA, 78(4): 24-37 \u222b with the United States Department of Agriculture that Wilson would work in partnership with Frank Meyer, the department's collector in China. Meyer, whose main interest was in plants of agricultural value, would also collect ornamentals in northern China, and Wilson would collect useful plants for the department in the southern zone. But Sargent was bitterly disappointed by how few ornamental specimens Meyer sent from Shanxi Province and was furious when these specimens were discovered to include several previously unknown species of larch (Larix), spruce (Picea), and pine (Pinus) from which Meyer, who had not recognized them as novelties, had not collected seed.5 Wilson, by contrast, was spectacularly successful, sending back thousands of herbarium specimens and large quantities of plant material, in the process enhancing the reputation of the Arboretum. Sargent, a man of strong opinions and personal self-confidence verging on arrogance, refused to accept Meyer's explanation that the north of China was \"an utterly barren region\"6 when it came to new ornamental woody plants and wanted to send a collector there to prove the contrary. Sargent also wanted this collector to harvest the botanical riches he was convinced were to be found in the high mountains of Shaanxi and Gansu Provinces in northwest- 26 Arnoldia 78\/4 \u2022 May 2021 Frank Meyer photographed larches (Larix gmelinii var. principis-rupprechtii) near Wutaishan, in Shanxi Province, in February 1908. Charles Sprague Sargent, suspecting these and other conifers in the region to be unique, wanted Purdom to revisit the site. ARNOLD ARBORETUM ARCHIVES cized (in Britain) record as a trade union activist\u2014 about which both Prain and Veitch appear to have maintained a discreet silence vis-\u00e0-vis Sargent\u2014meant that most potential employers saw him as a troublemaker, a label which would have made it very difficult for him to find employment in Britain. The first few weeks of 1909 passed in a blur, as Harry Veitch organized detailed briefings for Purdom on China. Purdom's instructors included Sir Robert Hart, recently retired after forty-eight years in China as inspector general of China's Imperial Maritime Customs Service, and Augustine Henry, the distinguished dendrologist who had spent nineteen years in China working for the Customs Service. The Kew-based photographer E. J. Wallis gave Purdom lessons in using a sophisticated glassplate camera.9 Purdom sailed on the Oceanic from Southampton to New York on February 3 and reached Boston four days later. Sargent immediately formed a favorable impression of Purdom,10 and he spent Purdom's second day in Boston writing an eight-page memorandum of guidance about where, when, and what to collect in China. Sargent told Purdom that, on arrival in China, he should seek out Ernest Wilson in either Shanghai or Yichang (in western Hubei Province) 11 before proceeding to Beijing. From there, he was to continue 120 miles north to Chengde (then often known as Jehol) and still farther north to the old imperial hunting ground at Weichang. In a characteristic display of wishful thinking, Sargent asserted that since Weichang \"has never been covered by a botanist, it is not impossible that you will find many interesting and possibly entirely new plants.\" Purdom was to leave Weichang in August so as to be in the Wutai mountain range, 180 miles southwest of Beijing in Shanxi Province, in mid-September, in time for the seed-drop of the conifers: obviously, Sargent especially desired seed from the new spruce, larch, and pine of which Meyer had sent herbarium specimens. Once the seeds had been collected, which Sargent thought \"ought not to take very long,\" he hoped that Purdom would return, via Beijing, to Weichang\u2014a round William Purdom 27 \u222b ern China. Sargent believed that, because the plants from that region endure harsh winters in their home range, they would be better able to stand the New England and north European winters than those from farther south. (The logic is seductive, and such plants will indeed withstand bitterly cold winters, but they are very vulnerable to late spring frosts, having evolved in a climate where spring is a brief prelude to a hot summer, a short transition from extreme cold to baking heat.) Sargent asked Isaac Bayley Balfour, the regius keeper of the Royal Botanic Garden, Edinburgh, for advice in identifying a collector, and Balfour recommended George Forrest,7 who had, in the spring of 1907, returned from a very successful three-year plant-hunting expedition in Yunnan Province and whom Balfour knew wanted to return to China.8 Sargent suggested to his old friend Harry Veitch, whose family firm, James Veitch & Son, dominated the British horticultural trade, that they jointly engage Forrest and share the harvest he would send back from China. Harry Veitch was agreeable, but although Forrest came to London in September to meet Sargent and Veitch, he refused their offer. Forrest was not impressed with the salary offered by Sargent and was reluctant to collect outside Yunnan, where he believed, quite correctly, that much more remained to be discovered. Nor would he agree to travel to China in early 1909 because he wanted to be at home for the birth of his first child in April. Sargent had to return to Boston in October, leaving Veitch to find a collector. After two months during which Veitch failed to propose a candidate, Sargent wrote to him in early December reminding him of their agreement to send a collector in early 1909. After consulting Prain and the director of the Kew Arboretum, William Bean, Veitch offered Purdom the post at a salary of two hundred pounds a year plus expenses of four hundred pounds a year. Purdom asked for time to think about it before agreeing on January 7, 1909. Truth to tell, Purdom had little alternative but to accept Sargent and Veitch's offer; his contract at Kew had expired, and he knew that his well-publiBaoding Wutaishan Beijing Chengde (Jehol) Dolon Nor Zhuizishan Mudanshan Yan'an Luoyang Jon\u00ea (Honan) Minxian Lanzhou Taibaishan Xi'an Lotani Gubeikou Shanghai SHAANXI SHANXI HENAN SHANDONG INNER MONGOLIA OUTER MONGOLIA SICHUAN HUBEI ANHUI JIANGSU ZHEJIANG GANSU ZHILI WEICHANG QILIAN MOUNTAINS QIN MOUNTAINS SHENYANG A M D O KOKO NOR 100 miles 200 kilometers 28 Arnoldia 78\/4 \u2022 May 2021 Purdom spent his fi rst collecting season, in 1909, north and west of Beijing. His second year centered on Shaanxi Province. In the third year, he collected in Gansu Province and the Tibetan region of Amdo. trip of around six hundred miles\u2014to gather seeds and herbarium specimens there. The year 1910 was to be spent in Shaanxi Province, where Purdom was to seek \"the wild tree peony\" (Paeonia suffruticosa) before exploring the mountain range near Xi'an, the ancient former capital. This region is around fi ve hundred miles southwest of Beijing. Finally, the third and last year, 1911, was to be spent in Gansu Province, in the high mountains on the border with Tibet, over one thousand miles from Beijing. All this was spelled out by Sargent with admirable clarity, and he was equally clear about the principal object of the expedition, which was \"to investigate botanically unexplored territory [and] to increase the knowledge of the woody and other plants of the [Chinese] Empire.\" In pursuit of this last goal, Sargent expected Purdom to dry six sets of herbarium specimens for all woody plants, including specimens of the same species that might occur in different regions so as to show the extent of any variation. He also wanted Purdom to photograph \"as many trees as possible,\" including their fl owers and bark, and \"if time permits [\u2026] views of villages and other striking and interesting objects, as the world knows little of the appearance of those parts of China you are about to visit.\" These goals were not quite the same as those articulated by Harry Veitch, who had told Purdom \"the object of your mission [is] to collect seeds and plants of trees and shrubs, also any plants likely to have a commercial value, such as lilies,\" but there was sufficient overlap that Purdom felt he could satisfy both his sponsors. ARNOLD ARBORETUM AND GIS COMMUNITY 1911 SEASON 1910 SEASON 1909 SEASON William Purdom 29 Purdom must also have welcomed Sargent's brief acknowledgment that it might be impracticable to complete the ambitious itinerary he had sketched out in three collecting seasons and that Purdom might need, in the light of local advice or experience, to change it. Sargent had his legal adviser draw up a contract, which he and Purdom signed. This stipulated that \"all seeds of herbaceous, alpines and bulbous plants and all bulbs and other roots except those of woody plants\" collected by Purdom would be the property of the firm of James Veitch & Sons and would be sent directly to them from China. Collections of woody plants would be divided equally between Veitch and the Arnold Arboretum. Photographs and herbarium specimens would belong to the Arboretum. The Arboretum would pay his salary and expenses in January and July, after which Veitch would reimburse one-half of the total sum involved. Purdom spent a fortnight in Boston, mostly being taught how to prepare herbarium specimens. This involves pressing specimens of plants in blotting paper (also known as drying paper), including, as appropriate, the leaves, stems, flowers, fruit, and seeds. It is a long and laborious process, not least because of the need to change the absorbent paper every couple of days until the plants are thoroughly dried out. These specimens are subsequently mounted on cardstock with a note of the name of the plant, if known, the date and site of collection, and any details recorded by the collector that may be lost as a result of pressing and drying, such as color or scent. After his training in Boston, Purdom traveled by train to Vancouver, from where he sailed for China on the Empress of Japan. He arrived in Shanghai on March 16, 1909. Ernest Wilson had repeatedly made it clear that he would hold Sargent to their two-year contract and was not interested in extending it. Nonetheless, when Sargent wrote to him that he and Harry Veitch had engaged Purdom and hoped that Wilson would brief him before returning to London, Wilson expressed disappointment at being \"passed over.\" But he promised that he would do anything he could to help \"your new man,\"12 and his briefing of Purdom in Shanghai seems to have been reasonably cordial. What is, however, clear from Purdom's full account of his briefing from Wilson13 is that Wilson did not suggest to Purdom that it would be to his advantage to engage any of the eight trained Chinese collectors who had supported Wilson over the last three years. Their contract with Wilson would end as soon as they had finished packing the harvest of the last season's collecting for shipment to Sargent. If Purdom had hired some or all of them, he would have benefitted from their experience and expertise in, for example, preparing herbarium specimens rather than having to train collectors himself, starting from scratch. The men themselves would surely have welcomed the continuation of their employment. Wilson's reticence is all the more noteworthy when one recalls that when Wilson started on his first collecting expedition to China in 1899, he was briefed by Augustine Henry (who was leaving the country) and immediately thereafter hired Henry's entire team, who had been trained over the previous decade.14 But Purdom lacked the experience to suggest he might do the same thing, and Wilson, despite his promise to Sargent that he would do all he could to help Purdom, did not propose it. One wonders whether Wilson kept silent because he anticipated that he might return to China within the three-year period for which Purdom was contracted to collect for Sargent and Veitch. In fact, in June 1910, Wilson did return and promptly reconstituted his team of helpers. Obviously, this would have been impossible if the men had been in the field with Purdom. A less charitable alternative explanation is that Wilson was not especially keen to provide Purdom with assistants who might help Purdom challenge Wilson's burgeoning reputation as the greatest of the Western plant hunters active in China.15 Certainly, in later years, Wilson quite deliberately burnished his reputation, including by rewriting some of the history of his first two expeditions.16 Immediately on his arrival in Beijing, Purdom applied himself to learning Mandarin Chinese, a language that he mastered remark- \u222b Clockwise from top: In the spring of 1909, Purdom traveled north of Beijing and crossed the Great Wall at the gateway town of Gubeikou. He continued northward by river and spent the summer in the imperial hunting grounds of Weichang. Although the region was predominately treeless, Purdom documented pines (Pinus tabuliformis) among the scattered forests. That fall, he returned to Beijing and headed west to Wutaishan, where he photographed a collection of Khingan fir (Abies nephrolepis) near his tent. The year 1910 was spent primarily in Shaanxi Province. He sent the Arnold Arboretum few photographs that year, but one showed the landscape of Mudanshan, where there was no sign of the wild tree peony. ALL PHOTOS ARNOLD ARBORETUM ARCHIVES Clockwise from top: In 1911, Purdom collected primarily in Gansu Province and Amdo, an adjacent region of Tibet, where he photographed a temple perched above the Tao River at Jon\u00ea. Purdom took a considerable number of portraits of families and individuals in the region. He also documented the dramatic mountains near Jon\u00ea, which he labeled as the Peling Mountains. Before returning to England, Purdom collected seedlings of the Chinese horsechestnut (Aesculus chinensis) at a temple in Beijing's Western Hills. ALL PHOTOS ARNOLD ARBORETUM ARCHIVES 32 Arnoldia 78\/4 \u2022 May 2021 ably quickly. Unusually for a Westerner in China at this time, Purdom consistently treated local administrators and farmers in the areas where he collected as his social equals, among whom he sought to make friends. Partly as a result, he was allowed into areas of China foreign travelers were actively discouraged from visiting, not least for their own safety. Purdom spent the 1909 collecting season in northern China and Mongolia, including in Wutaishan. Sargent had specifically tasked Purdom with collecting seeds from spruce and larches found there, which were not in cultivation in the West, but the wet summer of 1909 meant that the trees did not set seed. Although Purdom sent cuttings and seedlings, Sargent complained that they had been poorly packed and that, as a result, many of them had died on the six-week journey to Boston.17 He was only partly mollified by seeds that were germinating in the Arboretum's greenhouses. In fact, Purdom had dispatched thirty parcels of seeds and bulbs from more than three hundred unique collections to Boston and London that year. These included rhododendrons and primulas, a fine blue anemone, several peonies, and three species of clematis, one of which, the downy clematis (Clematis macropetala), has particularly graceful deep blue bell-shaped flowers. It first flowered in Veitch's Coombe Wood nursery in 1912 and remains very popular today. For Sargent, there were several poplars (Populus), elms (Ulmus), larch, and herbarium specimens of a new form of bird cherry (later named Prunus padus var. pubescens forma purdomii), which is a small tree with copious white racemes, bright red berries, and fine foliage. In April 1910, after overwintering in Beijing, Purdom traveled to western China. Sargent had asked him to investigate Moutan-shan (or Mudanshan, which translates to \"peony mountain\") near the ancient city of Xi'an, where he hoped Purdom would find the original wild peony. When Purdom arrived, however, he found that the plants had long ago been harvested for traditional medicines and the mountain was stripped bare. Purdom took several photos of the mountain to leave Sargent in no possible doubt that there were no peonies (and few other plants) there. Purdom had better luck near Yan'an, where he found a wild population of the tree peony. He ultimately collected over five hundred seeds of this dark red peony, which was raised in both Boston and Coombe Wood. (Sargent would later write of this as a \"first-rate achievement.\"18) On Taibaishan, in southern Shaanxi, he found a fine rhododendron with dark pink buds shading into white flowers, subsequently named Rhododendron purdomii. He also found another wild population of the tree peony, but with no seed. The next year, Purdom continued westward to Gansu Province and the Amdo region of Tibet. He found, in a monastery garden, a lovely winter-flowering viburnum (Viburnum farreri, then known as V. fragrans). He sent seeds to Veitch, who grew them on and subsequently sold his stock to Gerald Loder, the owner of Wakehurst Place in Sussex, where, in 1920, they flowered for the first time in Britain. Purdom also sent seed of an edible honeysuckle, Lonicera caerula, whose curious cylindrical fruit is today sold in the West as \"honeyberry.\" He ended the season in Minxian, in Gansu Province, where he had no choice but to wait for order to be restored following the anarchic violence that followed the Xinhai Revolution in October. Fortunately, Purdom had more or less completed the season's collecting, which included several fine primulas and asters, and in December, he was able to persuade the Minxian authorities to provide (for a fee) an armed escort to enable him to return, via Honan, to Beijing. When Purdom told the political staff at the British Legation about the attempted ambush near Shenchow, they were horrified to hear that he had killed three of the attackers, whom they strongly suspected (or they may have had confidential information confirming it as a fact) had been off-duty government soldiers.19 They urged Purdom not to repeat the story to anyone else lest he (and, by association, Britain) should be seen as taking up arms against the Chinese government. This advice suited Purdom, a very private man who throughout his life avoided personal publicity. Furthermore, Purdom was angling for a job with the Chinese Republican William Purdom 33 Purdom and two assistants make their camp on or near Mudanshan, in May 1910. His herbarium presses are arranged in the foreground, with his lever-action rifle resting against the central press. government and may well have believed that to publicize the shooting wouldn't help his prospects. He did give Sargent and Harry Veitch very brief accounts of the incident,20 but it was not reported in either the Chinese or English press, nor did he ever allude to it in later life. Both sponsors of the expedition were disappointed by Purdom's harvest. Harry Veitch recognized that \"if the plants were not there, then he [Purdom] could not send them,\" but Sargent was reluctant to accept that while his decision to send Purdom to the botanical terra incognita of northwestern China had been a perfectly reasonable throw of the dice, the gamble had failed. That would have meant recognizing that Sargent had got it wrong, and he chose instead to blame Purdom for not trying hard enough.21 Sargent also rebuffed Purdom's request to return home from Beijing via San Francisco and New York in order to enable him to visit Boston to explain why the results of the expedition had not matched Sargent's over-ambitious hopes.22 And the statistics that Sargent reported in his 1910-11 Annual Report to the President of Harvard University tended (at least) to leave readers with the impression that Purdom's harvest over the 1910 season had been less than one-quarter of Wilson's, whereas, in fact, he had sent the Arboretum and Veitch germplasm from almost \u222b \u00a9 BRITISH LIBRARY BOARD 34 Arnoldia 78\/4 \u2022 May 2021 exactly half the number of different plants collected by Wilson in the same season.23 Sargent's harsh judgment of Purdom's competence as a collector may well have been influenced by his comparing Purdom's collections with those of Ernest Wilson, sent from Sichuan Province. Such a comparison would prima facie not be to Purdom's advantage: the two men were not competing on a level playing field. The climate of Sichuan is subtropical, shading into tropical, and the annual monsoon delivers plentiful rainfall. Gansu, Shanxi, and Shaanxi Provinces, where Sargent had dispatched Purdom, share a temperate climate, with bitterly cold winters and little rainfall. Unsurprisingly, the flora of Gansu and its immediate neighbors is much sparser than the vegetation of Sichuan where Wilson principally collected. The Hengduan Mountains in western Sichuan illustrate the extreme biodiversity of the region where Wilson was collecting. The mountains are far enough south that during the last ice age they escaped being scraped bare by glaciers. The substantial variation in altitude created a range of habitats, from river valleys to alpine meadows and peaks, and a huge range of plants flourished there while those further north were wiped out by the ice. In consequence, the Hengduan massif is a biodiversity hotspot, a veritable plantsman's paradise in which it is estimated there are over 8,500 species of plants, 15 percent of them endemic (found only in that confined geographical area). They include over one in four of the world's species of rhododendrons (224 species), primulas (113 species), and mountain ash (Sorbus, 36 species)\u2014the list goes on and on.24 In contrast, plant biodiversity where Purdom was collecting was much lower. In the Qilian Mountains of Gansu, researchers have tabulated around 1,044 species of plants, and in southeastern Gansu, the number is around 2,458 species.25 Neither Wilson nor Purdom ever claimed to have done more than explore part of the provinces in which they hunted for plants, but the bottom line is that Wilson was collecting in a region where there was approximately three and a half to eight times the number of plant species than in the area to which Purdom had been sent by Sargent and Veitch. This made it almost inevitable that Wilson would send back to Boston specimens and seeds of more species than Purdom. In 1910 and early 1911, the only season for which it is possible to make a direct comparison, Purdom sent back to Harry Veitch germplasm associated with 374 unique collections numbers, while Wilson sent back 744 collections, 271 of them collected by his assistants after he had broken his leg.26 Sargent's negativity towards Purdom may also have been influenced by his feeling a measure of responsibility towards Wilson in respect of the avalanche that had nearly caused him to lose a leg and that left him with a severe limp.27 Wilson hadn't really wanted to go on the expedition, but Sargent had effectively forced him to, and it seems quite possible that he subconsciously vented a feeling of guilt about what had befallen Wilson on Purdom. Furthermore, the extent to which Wilson's work in China captured the imagination of the United States media and public meant that Wilson found a ready market for the articles and books that Sargent encouraged him to write about his expeditions. Wilson stressed his links with the Arboretum in the publications, and his star status, in turn, added luster to the fundraising efforts in which Sargent was constantly engaged to support the Arboretum and its activities. In short, it suited both men very well for Wilson to be front and center of the public stage, and there is nothing to suggest that either of them was concerned that the accomplishments of other collectors, including Meyer and Purdom, were overshadowed as a result. The final blow to any hopes Purdom entertained that this expedition might allow him to forge a reputation among the horticultural cognoscenti that would help him to secure a good job in Britain or the United States fell on his return to England. Harry Veitch had decided to close the firm, which had dominated the English nursery trade for decades, and sell the stock at auction, causing Purdom's collections to be dispersed and brought to market without his name being associated with them (Viburnum farreri, mentioned above, is a particularly egregious example). Purdom (left) returned to China in 1914 and spent two years collecting with the British botanist Reginald Farrer. Purdom used a clockwork self-timer to photograph himself with Farrer (right) and Zhang Bing Hua, the viceroy of Koko Nor (present-day Qinghai Province). This is the only known photograph of Purdom and Farrer together. \u00a9 BRITISH LIBRARY BOARD 36 Arnoldia 78\/4 \u2022 May 2021 All things considered, if we factor in Purdom's fundamental modesty and aversion to publicity, it's easy to see why he never captured the public imagination in the way that, say, Wilson or Forrest did. In 1912, Purdom began corresponding with officials in Beijing about a possible post in a yet-to-be-formed Chinese Forest Service, which would enable him to pursue an objective to which he was personally and strongly committed, namely the reforestation of China after decades of extensive and largely uncontrolled logging. There were long bureaucratic delays in setting up the service, and in 1913, when the alpine plant expert and plant hunter Reginald Farrer invited Purdom to join him on an expedition to northwestern China and Amdo, he accepted.28 He and Farrer botanized successfully in 1914 and 1915, collecting inter alios some fine poppies, alpines, primulas, and an elegant butterflybush (Buddleia alternifolia). Although Farrer would go on to write two of the best travel books of the era about the expedition, 29 the devastating effect on European gardening and horticulture of the First World War and the complete collapse in demand for new plants brought an abrupt end to their plant hunting at the close of 1915. In the spring of 1916, the Chinese government at last formally created a Chinese Forest Service, and Purdom was appointed as a senior forestry adviser to the Chinese government. Purdom must have been deeply happy at last to have achieved a senior management position in which he could make his mark. He began working with Han Ngen (Han An), the secretary of the Ministry of Agriculture, to train Chinese foresters, develop tree nurseries, and plant trees where they would do the most good. By 1919, after three years of backbreaking effort, over one thousand tree nurseries had been established in China, containing one hundred million young trees. In the same, year twenty to thirty million trees were planted on over one hundred thousand acres of otherwise unproductive land.30 Many of these were timber trees new to China, mostly from North America, which Purdom knew would do well in different Chinese regions and climatic zones. He organized the importation of many millions of seeds and cuttings, making him the only Western plant hunter to have imported into China vastly more plant material than he ever collected there. It appears that eventually Purdom and Sargent were reconciled: in 1920 and early 1921, Purdom is known to have sent plant material to the Arnold Arboretum. Frustratingly, however, there is no surviving correspondence from this time in the Arnold Arboretum files, and Sargent's personal papers are lost. Purdom died suddenly in Beijing in November 1921 at the age of forty-one, due to an infection contracted following a minor surgery. He was buried in the English cemetery in Beijing, but fifty-four of his Chinese friends and colleagues clubbed together to commission a large and elegant memorial stele in the Forest Service plantation at Xinyang, which they renamed the Purdom Forest Park. Remarkably, the stele and the park were both left alone during the violently anti-foreigner Cultural Revolution of the 1960s and 1970s and they are both carefully preserved to this day. The epitaph is too long to quote in full, but a hundred years later the sorrow felt by Purdom's friends who subscribed to the stele is still very clear. Perhaps what would have most pleased Purdom is their description of him as \"a true and loyal friend of the Chinese people who won the admiration and respect of his colleagues, worked tirelessly for the reforestation of China and who, had he lived, would certainly have trained the next generation of Chinese foresters.\" Will Purdom was a fine and honorable man, who rose from a position of very limited personal agency and overcame formidable obstacles to leave the world a better place for his passage. Not only does he deserve to be remembered in his own right, but his life has a good deal to teach us about our place in this interconnected world. His concerns about protecting local ecosystems are a reminder that these ideas were current well over a hundred years ago. Finally, we should, in justice, remember him when we plant his introductions in our gardens: among them, \"his\" viburnum, butterflybush, or bird cherry. \u222b William Purdom 37 Endnotes 1 Purdom letter to Harry Veitch, 23 March 1912 (copied by Veitch to Charles S. Sargent, 10 April 1912), Arnold Arboretum Horticultural Library, Harvard University (AA archive). 2 Anon. 1921. William Purdom. Journal of the Arnold Arboretum, 3(1): 55-56. 3 David Prain letter to Harry Veitch, 31 December 1908, Royal Botanic Garden, Kew, archives. 4 Ernest H. Wilson letters to Sargent, 21 November 1908 and 9 March 1909; also Sargent letter to Veitch, 26 April 1909, AA archive. 5 Sargent letter to Wilson, 8 July 1908, AA archive. Sargent also expressed his disappointment to David Fairchild, Meyer's superior at the Department of Agriculture. 6 Frank Meyer letter to Wilson, 7 May 1907, AA archive. 7 Bayley Balfour letter to George Forrest, 26 August 1908, Royal Botanic Garden Edinburgh archive. 8 George Forrest (1873-1932) made a total of seven expeditions to China, in the course of which he collected over thirty thousand different plants and herbarium specimens, nearly all of them from Yunnan Province in southwestern China. 9 Purdom was an apt pupil, and the Arnold Arboretum archive has a large collection of his photographs, which are an important resource for our understanding of remote areas of China in the first decades of the last century. 10 Sargent letter to Veitch, 16 February 1909; and to Prain, 25 February 1909, AA archive. 11 Sargent letter to Purdom, 8 February 1909, AA archive. 12 Wilson letter to Sargent, 9 March 1909, AA archive. 13 Purdom letter to Sargent, 26 March 1909, AA archive. 14 For a full account and a photo of the team, see: O'Brian, S. A. 2011. In the footsteps of Augustine Henry (p. 68 et seq.). Garden Art Press. 15 Wilson's biographer, Roy W. Briggs, suggests that Wilson was concerned that his replacement by Purdom might be seen as an adverse reflection on the quality of his own work in China. 16 See, for instance: Holway, T. History or romance? Garden History, 46(1): 3-27. 17 Sargent letter to Purdom, 3 May 1910, AA archive. 18 Sargent letter to Veitch, 13 June 1912, AA Archive. 19 On March 10, 1912, the political department of the legation sent a telegram about the ambush to the Foreign Office in London, but unfortunately it has been \"weeded\" from the file in the Public Record Office. The legation also asked the representative of the London Times in Beijing, Ernest Morrison, not to report the incident, and Morrison complied. 20 In addition to the letter that Purdom sent to Harry Veitch cited above, see: Thomas, W. B. 1913, July 10. Creator of 2,000 new plants. Daily Mail, p. 3. 21 Frank N. Meyer letter to David Fairchild, 15 October 1912, USDA compilation of Fairchild correspondence held at the University of California, Davis, Vol. 3, pp. 1600-1601. 22 Meyer letter to Fairchild, 21 December 1912, USDA compilation, Vol. 3, pp. 1619-1621. 23 For my full accounting of this, see: Gordon, F. 2021 Will Purdom: Agitator, plant-hunter, forester (pp. 111- 116). Royal Botanical Garden Edinburgh. 24 See: Kelley, S. 2001. Plant hunting of the rooftop of the world. Arnoldia, 61(2): 2-13. These figures are likely to have changed slightly in the intervening twenty years as new species have been identified and others have been reclassified. By way of comparison, the British Isles presently (2021) have 1,443 species from 308 genera, only 1.2 percent of them endemic. 25 Wang, J, Che, K., and Yan, W. 1996. Analysis of the biodiversity in Qilian Mountains. Journal of Gansu Forestry Science and Technology; also, Lu, W-Z. and Ren, J-W. 2005. Plant biodiversity and its conservation in Maijishan Scenic Regions of Gansu. Journal of Northwestern Forestry University, 20(4): 44-47. 26 Plant collecting is emphatically not a \"numbers game,\" and it would be foolish to use these figures to attempt to compare the relative efficiency of the two men. But Purdom clearly did a good job in a poor collecting area. Again, for my accounting of these numbers in the biography, see pp. 111-116. 27 For a full account of the story surrounding Wilson's accident, see: Dosmann, M. 2020. A lily from the valley, Arnoldia 77(3): 14-25. 28 Purdom letter to Reginald J. Farrer, 9 September 1913, Royal Botanic Garden Edinburgh archive. 29 See Farrer's books On the Eaves of the World (1917) and The Rainbow Bridge (1921). Both books are dedicated to \"Bill\", i.e. Will Purdom. 30 Reisner, J. H. 1921. Progress of forestry in China 1919- 1920. Journal of Forestry, 19(4): 396. The map in this article was inspired by the map on page 72 of Will Purdom: Agitator, Plant-Hunter, Forester and was created using Esri, Airbus DS, USGS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, Garmin, GSA, Geoland, FEMA, Intermap and the GIS user community. Purdom Plants at the Arnold Arboretum As of this writing, visitors at the Arnold Arboretum can find twenty-five trees and shrubs that arrived directly from Purdom (as seed) or Veitch (as plants) from Purdom's first expedition to China. Another twenty-six plants represent other Purdom lineages, including Forsythia that originated from Purdom's collections with Reginald Farrer. To map them in the landscape, visit https:\/\/ arboretum.harvard.edu\/explorer\/. Use the advanced search and input \"Purdom\" in the collector field. Francois Gordon retired from the British Foreign Office in 2009 after thirty years mostly spent in Africa. Today, he lives and gardens with his wife Elaine in Kent. His first book, Will Purdom: Agitator, Plant-Hunter, Forester, was published by the Royal Botanical Garden Edinburgh in 2021. It can be purchased on Amazon."},{"has_event_date":0,"type":"arnoldia","title":"George Ware and the Thornhill Elm: A Vision of Trees for the Future","article_sequence":6,"start_page":38,"end_page":47,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25734","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25e816e.jpg","volume":78,"issue_number":4,"year":2021,"series":null,"season":null,"authors":"Shearer, Kim","article_content":"In 1987, a plant pathologist in Montana ended an incomplete experiment by cutting down fourteen young American elm trees (Ulmus americana). At the time, Dutch elm disease (Ophiostoma ulmi, DED) was taking hold in parts of Montana. The only management practices then available in Montana were tree removal or pesticide sprays to stop the movement of the vectors, elm bark beetles. The pathologist, Gary Strobel, had been hoping to develop an unconventional method of disease management\u2014vaccinate the tree with a genetically engineered bacterium (Pseudomonas syringae) to fight the fungal disease. In lab trials, Strobel and his colleague Donald Myers had demonstrated that Pseudomonas syringae produced natural antibiotics that suppressed the spread of DED through vascular tissue. Still, Strobel needed permission from the Environmental Protection Agency to proceed with a field experiment that involved a genetically engineered organism. The bureaucratic machinations promised to delay the field experiments for another year, so Strobel moved forward with his experiments and injected fourteen trees before receiving formal approval. This moment captured national headlines focusing attention on the debate over genetically engineered organisms and drawing unwelcome attention to Montana State University, where Strobel was a researcher. Rather than put his colleagues at risk of losing federal funding due to his decisions, Strobel volunteered to destroy his field experiment. As an unanticipated result, however, the news also generated a newfound interest in tree breeding work occurring in a Chicago suburb. At the Morton Arboretum in Lisle, Illinois, George Ware had been busy developing DEDresistant elms the conventional way, through targeted breeding efforts using disease-resistant germplasm. In a 1987 New York Times article titled \"Fighting Elm Disease the Natural Way,\" Ware is quoted as saying, \"Dr. Strobel was trying to help one kind of elm quickly. We're looking more toward diversity over the long run.\" This quote highlights the nature of Ware's Elm Improvement Program and the efforts that he went through in developing the next DEDresistant elms. George Ware arrived at the Morton Arboretum in 1968, a mere two years after Marion Trufant Hall was hired as director and charged to lead the arboretum in an initiative to expand our research capacity. Ware was enlisted as the research director and the in-house ecologist and dendrologist. In 1972, four years into his tenure, Ware began noticing a peculiar tree in the arboretum collections: a stately elm growing outside the former study of Joy Morton, the arboretum's namesake and founder. At the time, much of the landscape in and around Chicago had been devastated by Dutch elm disease\u2014as was the case across the United States. The graceful American elm had been widely planted in the Chicago area, along streets and in parks. The prevalence of this species, which was also a ubiquitous forest tree, would ultimately be its undoing, enabling the rapid spread of both the vector and the disease by air and by root-to-root transmission. The first detection of Dutch elm disease in the United States was recorded in 1929 by Curtis May, a plant pathologist for the United States Department of Agriculture (USDA). May received samples collected in Ohio by plant pathologist Paul Tilford. The trees in Ohio were dying and a cause for concern. Later, in 1933, a USDA inspection would discover the source of the introduced disease: shipments of imported burl logs harboring the European elm bark bee- Facing page: In 1972, George Ware observed an elm at the Morton Arboretum that displayed exceptional form and resistance to Dutch elm disease. The tree would become Ware's first commercial tree introduction: the Accolade elm (Ulmus davidiana 'Morton' Accolade\u2122). PHOTO: STERLING MORTON LIBRARY SHEARER, K. 2021. GEORGE WARE AND THE THORNHILL ELM: A VISION OF TREES FOR THE FUTURE. ARNOLDIA, 78(4): 38-47 \u222b George Ware and the Thornhill Elm: A Vision of Trees for the Future Kim Shearer tle (Scolytus scolytus). The larvae of elm bark beetles, including our native species (Hylurgopinus rufipes), feed on the vascular tissue of infected trees, picking up spores with their bodies. When they mature and emerge from the tree, they can move to uninfected trees, introducing fungal spores. Newspapers across the country began raising the alarm about the rapid loss of trees as the disease continued to spread in the East and Midwest. By 1970, the Chicago region was reported to have lost more than fifty thousand trees and was projected to lose another fifty thousand within two years. It was amidst this devastation, in 1972, that Ware noticed the tree growing outside of Joy Morton's study window at the Thornhill Estate. It was an elm (Morton accession 2352-24*1) with gracefully arching branches, healthy and green foliage, and no symptoms of the disease. The original elm, fondly referred to by Morton Arboretum staff as the Thornhill Elm, was accessioned into the collections in 1924, shortly after Joy Morton founded the arboretum on his estate in 1922. With guidance from Charles Sprague Sargent, the director of the Arnold Arboretum, Morton established a 735- acre arboretum that included an herbarium, library, and nurseries, along with staff to manage it all by the time of his death. The most integral component of the arboretum\u2014the living collections\u2014included many accessions of plants initially sourced from the Arnold. In the initial establishment of the Morton collections, Sargent provided access to seed, clonal propagation material, and plants. One such packet of seed was labeled Ulmus crassifolia (the cedar elm), and records indicated that the seed had been wild-collected in Brownwood, Texas, by botanist Ernest Jesse Palmer. It was accessioned into the arboretum collections, and seeds were germinated and grown in the nursery. Eventually, a sapling was planted outside the bay window of Morton's study. As the years passed, the tree witnessed Morton's family and guests enjoying summer afternoons by the pool. The sloping vista beneath the elm was crowned by hawthorns for which the estate was named. There were staff picnics for Morton Salt Tree breeding is a slow, steady process, requiring years to grow and evaluate each generation of hybrids. Over the decades, Ware's Elm Improvement Program would produce some of the most popular disease-resistant elms for the North American landscape. STERLING MORTON LIBRARY \u222b George Ware 41 Company and the Morton Arboretum in the coming decades. The tree overlooks the Morton family cemetery and bore witness to family funerals, but it also provided shade to guests at weddings and garden soirees. Eventually, when the crumbling mansion was demolished long after Morton had died, the tree stood guard over Morton's study, which was preserved as part of a new facility for educating the public about plants and the rest of the natural world. In 1972, Ware looked at this tree and recognized that it was, in fact, not Ulmus crassifolia. The leaves were too large, the bark not quite right, and the form much too refined. As a dendrologist who had been a faculty member at Northwestern State College in Louisiana, Ware was familiar with U. crassifolia, which is native to that region. In fact, one of the first deposits Ware made into the arboretum collections in 1968 was a packet of cedar elm seed (Morton accession 385-68) that he had collected from the wild in Seguin, Texas. After further investigation (and even a visit to Arnold), Ware confirmed that the Thornhill Elm was U. davidiana, a species native to eastern Asia. Noting the native origin of the species and the lack of symptoms in the tree, Ware saw the possibility that the Dutch elm disease pathogen had Asiatic origins itself. Perhaps the presence of Ophiostoma ulmi in the natural habitat of U. davidiana had led to coevolution of the species such that the David elm had adapted a natural biochemical defense mechanism to combat the disease. In this tree, Ware saw great potential. The Thornhill Elm inspired the development of the first breeding program at the Morton Arboretum, the Elm Improvement Program. As a trained ecologist and dendrologist, Ware understood the necessity for genetic diversity within a population. He was soon on the search for more parent material to include in his germplasm collection. By 1980, Ware had clones of the Thornhill Elm propagated and under evaluation. That same year, he published two articles in the Journal of Arboriculture focusing on the qualities necessary for trees to survive in human-built landscapes and the attributes of Asian elm species that made them ideal candidates for such an environment. These publications were an effort to raise awareness within a community of tree experts about the possibilities that were held within the genetic resources of Asian elms. While American elms were being felled across the eastern United States, Ware was proposing a new solution to a decades-old problem: Let's plant Asian elms, he suggested, given that these species are adapted to both the constructed environment and the Ware recognized the value of Asian elm species as urban trees in North America. This list outlines species Ware recommended for evaluation and breeding in the Journal of Arboriculture and Landscape Plant News. Distribution and descriptions have been adapted from his papers. Species Geographic distribution Ware Description Selections available in US nursery trade? Ulmus davidiana (syn. include U. japonica, U. wilsoniana, U. propinqua) China, Japan, Korea, Mongolia, Siberia Variation in habit; tolerant of hostile conditions Yes; many introductions made in the past couple of decades U. glaucescens Gansu Province (China), northern China Small tree; small leaves, fine texture; yellow to orange fall color; tolerant of urban conditions based on its distribution No U. laciniata Humid areas of northern China, Korea, Siberia, and Japan Small to medium tree; potential drought hardiness; lobed leaves; Zelkova-like branching No U. macrocarpa China, Mongolia, Korea, and Siberia Strong wood; shrub to mediumsized tree; adapted to humid and arid regions; tolerant of \"hostile\" conditions No U. parvifolia China, Korea, Japan Tolerant of drought, pollution, poor soils; attractive lace bark; glossy leaves Yes; many introductions made in the past couple of decades devastating disease. Clones of the Thornhill Elm are now widely available in the commercial nursery trade under the name Ulmus davidiana 'Morton' Accolade\u2122. When developing any plant breeding program, a breeder must first start with objectives and further refine them by identifying specific desirable traits. Ware's primary objective was to develop elm trees with Dutch elm disease resistance. Second to that, he aimed to develop trees that were not preferred by the elm bark beetles or elm flea weevils (Orchestes alni). Beyond pest and disease resistance, Ware would focus on species adapted to environments of the extremes: temperature, drought, flood, high winds, blizzard, and \"hostile\" soils. He defined hostile soils as those with high pH, poor aeration, and minimal organic matter. He noted that these are all common conditions of the Midwest and Great Plains, and coincidentally, these are the same conditions faced by trees in developed landscapes regardless of the region. Ware went on to list and describe Asian elm species that should be considered for breeding programs. Meanwhile, Ware began the process of hybridizing elms that were available within the Morton collections. He collected branches covered in rounded floral buds and brought them into his lab. He placed the cut stems in vessels containing water and positioned them upon white sheets of paper spaced out along lab benches. As the forced stems began to flower, yellow piles of pollen would accumulate on the paper, signaling the pollen was ready to be collected and stored. Ware then used a ladder to take this pollen into the canopy of a female parent tree, where he secured a bag over a flowering stem. Making an opening in the bag, he dispersed pollen inside and mimicked the movement of the wind to ensure the pollen made contact with the receptive stigma. Once the bag was securely shut, he climbed down from his ladder and waited. This process led to the development of several new hybrid elms, including Ulmus 'Morton Glossy' Triumph\u2122. This selection is one of the most popular of Ware's elm introductions due to its low maintenance requirements in both commercial nursery production and municipal tree management. While a breeder can develop the best possible plant selection, the plants would not get very far out of the breeding program without help from the nursery industry. Ware was acutely aware of this. While his initial collaborations were with arborists, foresters, and botanists, he would go on to develop strong working relationships with the nursery industry, specifically Keith Warren, the former manager of new plant development for J. Frank Schmidt & Son, based in Boring, Oregon. The two first discussed the possibility of evaluating Ware's elm selections after a Metropolitan Tree Improvement Alliance conference, hosted at Thornhill in June of 1990. This meeting would lead to a collaboration between the Morton and J. Frank Schmidt that continues today, enabling hybrid elm selections to be propagated on greater scales and evaluated in field research. The first grafting of Ware's elm hybrids at J. Frank Schmidt occurred in 1994\u2014just twenty-two years after Ware recognized the tree's potential and seventy years after being received as seed labeled Ulmus crassifolia. The Oregon Department of Agriculture helped the collaborators set up a screened isolation and quarantine area at the commercial nursery, ensuring that DED would not be introduced into the Oregon landscape due to the nursery trade. By 1995, additional propagation material was distributed for in vitro propagation evaluation by Microplant Nurseries, a tissue culture lab based in Gervais, Oregon, managed by Gayle Suttle. At that time, there were not yet any cultivars of U. davidiana available through the commercial industry. Ware also recognized that for elm breeding efforts to be effective, additional genetic material needed to be collected from the wild. When he began his research, he found that few elms of wild provenance were available in the collections of North American public gardens, potentially creating a genetic bottleneck for any North American elm breeding program. The total number of elm species is somewhere within the range of twenty to forty, depending on taxonomic classification, and the center for Facing page: To develop new elm hybrids, including Ulmus 'Morton Glossy' Triumph\u2122, Ware carefully crossed select trees using pollination bags, secured high within the tree canopy. PHOTO: JIM NACHEL, STERLING MORTON LIBRARY 42 Arnoldia 78\/4 \u2022 May 2021 \u222b this diversity is unmistakably in eastern Asia. The Flora of China indicates that more than half of all elm species are native to the region. Ware and his colleagues ultimately visited China five times and the Soviet Union three times, developing relationships with forestry researchers willing to collect seeds in the wild and ship them to Ware. Today, the Morton's elm collection contains 329 accessioned individuals representing thirty-three species and thirty-four cultivars, a dramatic increase from 1968, when Ware arrived. At that time, the elm collection included fifty-one trees, which represented nine species and ten cultivars. Of the newer individuals accessioned into the Morton collections, eighty-one came directly as plants from Ware's breeding and research program. Ware also actively distributed seed and plants throughout the United States. He coordinated a seedling distribution program through which he distributed one thousand seedlings to Midwestern nurseries, aiming to popularize the Asian elm species. Municipal foresters and park managers regularly arrived at the Morton Arboretum's service gate searching for elm seedlings he had promised. As the current manager of the program that Ware initiated, I still receive notes from recipients of such gifts who recount fond memories of Ware and his generosity. Today, the seedling trees that he distributed can be found from Oregon to New York and Illinois to Louisiana. Several of Ware's elms were even planted in the late 1980s on the course of the Winged Foot Golf Club, the prestigious host of multiple US Opens in Mamaroneck, New York. This planting was a direct result of a 1987 New York Times interview of Ware following the Strobel controversy. By 1990, Ware had several elm selections in the pipeline and a greatly expanded collection of germplasm. He then began the process of developing a new breeding population. Working with large, wind-pollinated, late-winterflowering trees presents unique challenges to a breeder. The flowers are insignificant and often located more than six feet above the ground. (I can attest to the complications of these factors as a breeder working with elms today.) To sim- This table outlines five of Ware's most well-known elm cultivars. Note that Ulmus japonica and U. wilsoniana are taxonomic varieties that make up the U. davidiana species complex, but they are listed here as the original species for the sake of simplicity. Information found in this table is adapted from the Chicagoland Grows' Plant Release Bulletin (no. 44). 44 Arnoldia 78\/4 \u2022 May 2021 Cultivar and trade name Parentage\/ origin Traits USDA Hardiness Zones Dimensions (feet) Ulmus 'Morton' Accolade\u2122 Chance seedling U. japonica x U. wilsoniana Vase-shaped habit and vigorous grower; foliage fine-textured, dark green, and glossy with yellow fall color; DED and elm yellows resistance; resistant to elm leaf beetle 5 - 8 20 year 30' H, 15' W Mature 50 - 60' H 30 - 40' W Ulmus 'Morton Plainsman' Vanguard\u2122 Chance seedling U. japonica x U. pumila Relatively upright branching and rounded habit in youth; requires corrective pruning to avoid included bark; dark green foliage with yellow fall color; DED and elm yellows resistant; susceptible to elm leaf beetle, Japanese beetle, and leafminer 5 - 7 Mature 45 - 50' H 40 - 50' W Ulmus 'Morton Glossy' Triumph\u2122 Controlled cross U. Accolade\u2122 x U. Vanguard\u2122 Grower favorite due to ease of training; lustrous dark green foliage with yellow fall color; upright oval form that ages to vase shape; strong branching; excellent DED resistance; moderate pest resistance 4 - 9 Mature 50 - 60' H 40 - 50' W Ulmus 'Morton Stalwart' Commendation\u2122 Controlled cross U. Accolade\u2122 x (U. pumila x U. carpinifolia) Symmetrical arching branches, upright oval habit; large, dark green leaves with yellow fall color; rapid growth and broad adaptability; excellent DED resistance; moderate susceptibility to elm leaf beetle, Japanese beetle, and gypsy moth (4)5 - 9 Mature 50 - 60' H 40 - 50' W Ulmus 'Morton Red Tip' Danada Charm\u2122 Chance seedling U. japonica Rounded habit in youth maturing to large and elegant vase-shape; fast grower; glossy green foliage with redpigmented new growth; yellow fall color; excellent resistance to DED and elm yellows; moderate susceptibility to Japanese beetle and elm leaf beetle (4)5 - 9 Mature 60 - 70' H 50 - 60' W plify the hybridization process, Ware developed an isolation block of sorts in a local cemetery. He knew the cemetery would not be paved and that the trees would be left alone until they declined from old age. While Ware retired in 1995, he continued to develop his vision of trees for the future as a research associate of Morton Arboretum until 2009. The selection criteria that Ware developed for this population include tolerance to DED and elm yellows (a phytoplasma disease, Candidatus Phytoplasma ulmi, which causes leaves to suddenly wilt in late summer), pest resistance, cold hardiness, vigor, and red fall color. Red foliage is not commonly seen in elms. Typically, the fall color is a muddy yellow. Ware, however, had noticed an intriguing trait in a group of Asian elm seedlings: red pigmentation in emerging leaves. He understood that if the seedlings could produce red pigmentation (anthocyanins) in leaves during the spring, they should be able to use the same biochemical pathway to produce anthocyanins in the fall. This unexpected discovery led to red fall color becoming a new breeding objective. I was hired as the tree and shrub breeder for the Morton Arboretum in 2016. When I arrived here, I was certainly not an elm expert. I had spent my graduate school years working primarily with shrubs and herbaceous perennials. It took some time to unearth the details of the Elm Improvement Program, but today, I can say that we are continuing to make progress with Ware's legacy project. The program is now part of the Daniel P. Haerther New Plant Development Program, named in honor of a generous benefactor of the arboretum who was one of many that Ware inspired to appreciate the development of trees for the urban landscape. Ware consulted about elms on Haerther's estate, and in the process, the two would develop a relationship centered on a love of trees. Currently, we have sixty-one seedling selections from the breeding population that Ware left behind for the next generation. These were all selected for fall colors ranging from oranges to reds and purples. The breeding population includes the Ulmus davidiana complex, a variable group that was historically treated as three STERLING MORTON LIBRARY Ware (center) embarked on plant-collecting expeditions to acquire new elm germplasm from populations in China and the Soviet Union. His collaborators on this 1990 expedition to Shaanxi Province, China, included (from left) Ross Clark, Peter van der Linden, Kris Bachtell, and William Hess. \u222b separate species (U. japonica, U. wilsoniana, and U. propinqua). The population also includes three other Asian species (U. macrocarpa, U. parvifolia, U. pumila), an Asian hybrid (U. 'Sapporo Autumn Gold'), and the European field elm (U. minor). Our primary focus has been on the U. davidiana complex. We have selected a tree that will serve as the seed parent. It has an attractive form and relatively petite stature, along with somewhat glossy and predation-free foliage in the summer. We also continue to expand the program, particularly with work on the lacebark elm (Ulmus parvifolia). In 1996, Ware published two short articles in Landscape Plant News regarding this Asian species. He had participated in a USDAsponsored research exchange trip to China led by Eugene Smalley from the University of Wisconsin-Madison. Ware and four American colleagues joined Smalley in the fi eld to collect seed and determine the natural range of U. parvifolia. They were also very much focused on building relationships with Chinese researchers. Ware considered this elm species to be an especially promising selection for built environments of the South due to its broad adaptability to heat, fl ooding and drought, hostile soils, and both humid and arid conditions. However, he also noted that it would not perform well in northern states due to limited cold hardiness. Today, lacebark elms have demonstrated some hardiness with proper site selection. At the Morton Arboretum, seventeen individuals of this species (along with three cultivars and three unnamed hybrids) have survived multiple polar vortexes. I have also witnessed the lacebark elm growing and thriving from North Carolina to New York City and Las Vegas to New Orleans. This widespread adaptability, however, is accompanied by legitimate concerns about weediness. Even though the lacebark elm has not been widely planted in the Midwest, it is already listed as a weed of concern in Wisconsin. Colleagues at public gardens in other regions have expressed similar apprehensions about the species. This concern has led us to develop a new elm improvement project at the Morton Arboretum focused on developing selections with reduced fertility. Breeders have long used methods of mutation breeding to develop seedless plants. The most commonly known examples include the seedless watermelon and banana. These were developed through a traditional breeding method referred to as interploidy hybridization. Ploidy is the number of complete sets of chromosomes found in the cells of an organism. Humans typically carry two sets of chromosomes (diploid)\u2014 STERLING MORTON LIBRARY A young Accolade elm represents the success of Ware's vision for tree breeding and introduction. This commercial introduction is now one of more than twenty-six cultivars of Asian elms available in North America. one set inherited from our mother, the other from our father. A plant, however, can carry many more sets of chromosomes within its cells. Having three sets of chromosomes (triploid) often causes issues in reproduction due to the odd number of chromosomes that cannot segregate evenly during meiosis. To develop a triploid, a breeder must hybridize a diploid and a tetraploid (four sets of chromosomes). Tetraploids can be developed through the application of chemical mutagens known as mitotic spindle fiber inhibitors. (One such chemical is colchicine, a toxic compound found in the autumn crocus, Colchicum autumnale.) We currently have several tetraploid lacebark elms, but now we must wait for them to mature. Once these trees have reached maturity, we will hybridize them with diploids in our collections that are reasonably cold hardy. Meanwhile, from the Ulmus crassifolia seed that Ware deposited in the Morton Arboretum collections in 1968, we have selected a tree with a remarkably symmetrical and pyramidal form that has survived severe winters and flooding events unscathed. We are building numbers of rooted cuttings to grow in evaluation blocks, inoculate with DED, and distribute to partners for evaluation around the country. Additionally, both this species and the lacebark elm are fallflowering species. Considering they are windpollinated and not self-compatible, we have begun collecting open-pollinated seed from our cedar elm selection and an adjacent lacebark elm, and we plan to evaluate the resulting seedlings. According to a paper published by USDA researcher Frank Santamour in 1973, not only are the two species compatible, but the lacebark elm confers increased DED resistance to its hybrid progeny. As Ware noted in his 1987 New York Times interview, the Morton Arboretum's effort to develop new trees for the American landscape has been focused on traditional breeding efforts. These slow and steady methods have required several decades, spanning multiple careers. Yet, the value in Ware's approach to breeding and outreach is evident in today's nursery catalogs and landscape. Once there were monocultures of American elm planted across the country in the built landscapes of cities and suburbs, but today the monocultures have been replaced with DED-resistant Asian elms. This diversity includes more than thirteen cultivars of elms from the Ulmus davidiana complex, in addition to at least thirteen cultivars of U. parvifolia. Many more selections of various species are still in the pipeline from academic and commercial breeding programs around the country. Through tenacity and vision, George Ware managed to inspire the nursery industry to adopt a new crop and introduce an unfamiliar Asian elm species to the North American landscape. The work has resulted in further diversification of our tree palette. It all began with the original Thornhill Elm, distributed to the Morton Arboretum by the Arnold Arboretum almost a century ago. Now, this very selection graces the landscape of the city of Boston, having come full circle in its journey from seed to cultivated tree. References Chicagoland Grows. 2017. The Morton Arboretum Elms. Plant Release Bulletin (no. 44). http:\/\/ www.chicagolandgrows.org\/downloads\/ MortonArboretumElms.pdf King, S. S. 1971, September 7. Dutch elm disease spreads westward. New York Times, 45. Malcolm, A. H. 1987, November 1. Fighting elm disease the natural way. New York Times, 9. Myers, D. F. and G. A. Strobel. 1983. Pseudomonas syringae as a microbial antagonist of Ceratocystis ulmi in the apoplast of American elm. Transactions of the British Mycological Society, 80(3): 389-394. https:\/\/doi.org\/10.1016\/ S0007-1536(83)80034-5 Santamour, F. S., Jr. 1973. Resistance to Dutch elm disease in Chinese elm hybrids. Plant Disease Report 57(12): 997-999. Schneider, K. 1987, September 4. Tearful scientist halts gene test. New York Times, A1, A11. Ware, G. H. 1980a. Little-known Asian elms: Urban tree possibilities. Journal of Arboriculture, 6: 197-199. Ware, G. H. 1980b. In search of new kinds of elms. Journal of Arboriculture, 6: 233-237. Ware, G. H. 1996a. Notes on elms observed on a trip to China. Landscape Plant News, 7(10): 4-6. Ware, G. H. 1996b. New elms for urban landscapes. Landscape Plant News, 7(10): 6-8. Kim Shearer is the tree and shrub breeder at the Morton Arboretum, where she is also manager of the Daniel P. Haerther Charitable Trust New Plant Development Program and woody plant liaison for Chicagoland Grows. George Ware 47 \u222b"},{"has_event_date":0,"type":"arnoldia","title":"A Temperate Cousin: Leitneria floridana","article_sequence":7,"start_page":48,"end_page":49,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25735","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25e8526.jpg","volume":78,"issue_number":4,"year":2021,"series":null,"season":null,"authors":"Enzenbacher, Tiffany","article_content":"\"Stop! We're here!\" directed Kea Woodruff, who was navigating from the passenger seat of our rented vehicle. Woodruff was then the Arnold Arboretum's plant growth facilities manager. We were on day three of a 2018 plant-collecting expedition to Arkansas and Oklahoma\u2014part of the Arboretum's Campaign for the Living Collections\u2014and we were driving up Highway 62 in northeastern Arkansas, approaching the Missouri border. Months prior, we reached out for guidance on our Ozark-specific taxa list to the Arkansas National Heritage Commission. They provided an account of a particular population of corkwood (Leitneria floridana), a rare shrub sparsely endemic to the southeastern United States. We dropped a Google pin on the approximate location of their 2003 description and hoped that no habitat loss occurred between then and October 2018. I steered onto the shoulder, and we began scouring the nearby flora as traffic whizzed by. After what seemed like only a moment, Woodruff pointed to a promising-looking stand. \"Wait,\" she inquired, \"isn't that it?\" Corkwood is a striking plant, and we were able to confirm it in short order. It is monotypic (the only species in its genus) and is in a mostly tropical family, Simaroubaceae. The most well-known and recognizable temperate member is the tree of heaven (Ailanthus altissima), a noxious urban weed. The noninvasive, but just as conspicuous, corkwood typically grows five to ten feet tall\u2014although it can reach up to twenty feet. It is adorned with elliptic olive-green leaves that are glossy, leathery, and crowded toward branch tips. The common name derives from the buoyancy of the wood. It is one of the lightest woods known and has been used to float fishing nets. The bark is a deep reddish-brown with lenticels. Corkwood is content to sucker and form thickets, particularly in its ideal environment: forested swamps and flooded soils. The Arboretum's inaugural corkwood plants (accession 5336) arrived from botanist Benjamin Franklin Bush, who sent plants in 1894, just two years after he had first documented the species in southeastern Missouri. (The species had been named, in 1860, by Alvan Wentworth Chapman, based on populations in the estuary of Florida's Apalachicola River.) The plants prospered along Meadow Road, in a location affectionately known as \"Leitneria swamp,\" where water accumulates and persists throughout most of the year. Eventually, this accession became indistinguishable from other corkwoods that were planted around 1970, and the mixed planting was given a new accession number (244-97). Plants from Taylor County, Florida, were later added to the location (accessions 29-96 and 30-96). The species is near threatened in the wild and remains in several Florida and Texas counties abutting the Gulf, and a few inland ones in Arkansas, Georgia, Mississippi, and Missouri. Woodruff and I were determined not to let several feet of muck act as a deterrent. I repurposed two herbarium voucher bags as waders and bounded in. As we had anticipated, we found no fruit. The olive-sized brown drupes are borne in clusters of two to four near branch tips, below the foliage. Corkwood is dioecious, having separate male and female plants. Flowers are axillary catkins approximately one-anda- half inches long. This colony could have been a single sex, or perhaps voracious critters beat us to the fruit. Plan B consisted of combing for small suckers. We dug three, which we bagged and labeled as puzzled drivers drove past. From there, we continued to the nearest FedEx location, where we were grateful for the kindness of strangers. After we explained that we hailed from Boston and were in the midst of an expedition, the store clerks were keen to facilitate packaging in the tallest boxes they had available. The plants arrived the next day at the Dana Greenhouse, where they were potted and catalogued (accession 278-2018). In April 2020, the three individuals were planted in the seep on Bussey Hill\u2014this time a location distinct from any others. I hope these plants will colonize the seep over the next decade, just as the original collection has done to the Leitneria swamp. Tiffany Enzenbacher is the manager of plant production at the Arnold Arboretum. A Temperate Cousin: Leitneria floridana Tiffany Enzenbacher"},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25698","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270ab6e.jpg","title":"2021-78-4","volume":78,"issue_number":4,"year":2021,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Building a Comprehensive Plant Collection","article_sequence":1,"start_page":2,"end_page":4,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25720","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24e816c.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Carstens, Jeffrey D.","article_content":"Building a germplasm collection can take years or, more realistically, even multiple careers to assemble. The United States National Plant Germplasm System has nineteen stations around the country, and the goal is to acquire, conserve, evaluate, and distribute genetically diverse plant material. As a genebank curator at the North Central Regional Plant Introduction Station in Ames, Iowa, I'm responsible for managing collections of woody plants like ashes (Fraxinus) and chokeberries (Aronia), and also herbaceous plants. The collections arise from plant exploration by staff members (I typically make at least five collection trips per year), through exchange with other genebanks or public gardens, or by specific arrangements between a curator and an independent collector. The latter became more important than ever in 2020, as the coronavirus pandemic restrictions prevented normal travel. One of our most notable collections from this unusual season occurred in the mountains of northeastern Tennessee. The story, however, began in June of 2018, when I sent an email to Roger McCoy, the director of the Tennessee Division of Natural Areas, looking for contacts in eastern Tennessee who might be able and willing to collect native Monarda species. Monarda, or the bee balms, is a group of herbaceous plants native to North America and Mexico and is represented by approximately eighteen species. Our Monarda germplasm collection in Ames currently includes fourteen species, represented by 164 accessions. In the last couple of years, we've acquired interesting samples, including three species that were first described by botanists within the past decade: M. luteola, found in northeastern Texas and southwestern Arkansas; M. austroappalachiana, endemic to the Southern Appalachians; and M. brevis, a dwarf, earlyflowering species found in West Virginia and historically in Virginia. McCoy connected me with Marty Silver, a park ranger at Warriors' Path State Park, who graciously volunteered to help. Silver stated he had \"limited botanical skills\" and was simply an \"interested amateur botanist spending spare time in the field in various wild places in Tennessee.\" To ensure initial success, we selected Monarda didyma as the target from eastern Tennessee, since we had no holdings of the species from the region. The species also displays very conspicuous red flowers from July through August and is somewhat ubiquitous in the target area. This would make the plants relatively easy to locate. By the end of August 2018, Silver had documented several flowering patches of M. didyma, and that fall, he returned and successfully collected seed (accession Ames 34356). Despite living approximately an hour away from the sampling site, Silver conducted this travel and exploration on a volunteer basis. As Silver and I communicated after the 2018 collection, he drew my attention to a very thorough floristic survey of the nearby Rocky Fork Tract, written by Foster Levy and Elaine Walker, published in 2016. Silver connected me with Levy, who brought our attention to several Monarda specimens from the area that were labeled M. x media, a taxon that was missing within our germplasm collection. We designated this hybrid as our next target. Monarda x media is of potential interest for development as an ornamental landscape plant. Moreover, when I reviewed the published literature and herbarium specimens, I found a curious backstory for the taxon, suggesting that welldocumented wild collections could also support taxonomic research. The taxon was described over two hundred years ago, in 1809, by the German botanist Carl Ludwig Willdenow, who published the name without the multiplication symbol. The symbol is used to indicate plants of hybrid origin, although it is not required in Building a Comprehensive Plant Collection Jeffrey D. Carstens Facing page: Collaboration is key for developing a plant germplasm collection. A recent seed collection of Monarda x media in northeastern Tennessee is a case in point. PLANT PHOTOS BY MARTY SILVER; SEEDS BY ASHLEY SONNER, USDA ARS NCRPIS Monarda 3 4 Arnoldia 78\/3 \u2022 February 2021 a taxonomic name nor does authorship change in the event a name is later recognized as a hybrid. While Willdenow's description does not suggest that he recognized this taxon as a hybrid, he nonetheless noted an affinity to M. fistulosa, commonly known as wild bergamot. By 1901, Merritt Fernald, a botanist at Harvard, described observing numerous intermediate forms of M. media, making separation from M. fistulosa difficult. Currently, Monarda x media is recognized as a variable group of plants with intermediate characteristics of M. didyma and either M. fistulosa or M. clinopodia or both. These numerous intermediate forms may stem from the various hybrid combinations, and thus, the name M. x media should ultimately be assigned to a specific combination (for instance, M. didyma crossed with M. clinopodia), with new names given to each of the others. Surprisingly, Willdenow did not designate a type herbarium specimen, which could make it more difficult to determine which combination should, in fact, retain the original name. To correctly sample true-to-type specimens of Monarda x media in nature, Silver would need to mark populations in bloom, since M. clinopodia\u2014a white-flowered species\u2014and M. didyma are often found nearby; sometimes they are even intermixed with M. x media. This raises an interesting question about whether M. x media plants are stable in nature or whether they require the parents to constantly resupply them. Despite subsequent discussion about conducting reconnaissance and sampling for M. x media in 2019, Silver had other projects that left no time to acquire samples. The following year, as implications of the coronavirus pandemic were becoming clear, I followed up by asking about the possibility of sampling a Monarda x media population. Silver quickly replied, \"I am much more out and about in the field (outside and distanced) these days. If pointed in the right direction, I'll be glad to try and find populations within my limited taxonomic skills.\" While the pandemic quickly resulted in travel cancellations and restrictions (out of state, not to mention out of the country) across many agencies, Silver saw being in the field as an opportunity to be completely distanced while regaining a sense of normalcy. Using Levy's herbarium vouchers, we identified a total of three potential sites, but since the specimens were described from a broad geographic area, their relocation was going to be challenging. A few weeks later, Silver reported finding Monarda x media while on a hike on his day off. His hike to get to these populations was three and a half miles (one way) with an elevation climb of over two thousand feet. He took notes, GPS coordinates, and photos. Making the hike once again in the fall, Silver relocated the five previously flagged flowering patches, but one patch had been completely destroyed and another patch was nearly decimated due to human disturbance. He collected seeds from the available patches and then shipped them to Iowa. I assigned them an accession number (Ames 35579) and deposited them into the repository's freezer, which maintains the seeds at 0\u00b0F (-18\u00b0C). This collection will be periodically monitored for viability, and when germination falls below a critical level, it will be regenerated using controlled pollination techniques ensuring the preservation of the genetic profile for the future. Since Silver sampled each clonal patch separately along with appropriate plant descriptions, the collections will be important resources for future research (including ecogeographic and phylogenetic studies). The collections might also be useful for selecting superior genotypes for the nursery industry. Having one collection of this taxon is, of course, only a start\u2014additional samples are desired. Yet Silver's collections demonstrate the critical importance of local assistance while assembling a comprehensive germplasm collection, especially given the amount of time and effort required to acquire even a single collection. In the end, I'll never forget Silver humbly labeling himself as an \"amateur botanist with limited taxonomic skills,\" as his Monarda x media collection is one of the most exciting, well-documented samples of Monarda that I've accessioned in my nearly twenty-year career. Jeffrey D. Carstens is the curator for woody and herbaceous plants at the North Central Regional Plant Introduction Station."},{"has_event_date":0,"type":"arnoldia","title":"A Conservation SOS: Polygonum hickmanii","article_sequence":2,"start_page":5,"end_page":6,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25721","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24e856f.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Forbes, Holly","article_content":"Plants with less-than-showy flowers tend to get overlooked, even by some of the sharpest botanists. When a plant is only a few centimeters tall and flowers later in the season than its more eye-catching neighbors, it can be even easier to miss. The Scotts Valley polygonum (Polygonum hickmanii) is a case in point. This tiny species was first described in 1995 and was already very rare. It occurs in a limited urban area in Scotts Valley, near Santa Cruz, California, where it is under pressure from development. Only 2,100 plants were observed in 1997, and in 2003, the United States Fish and Wildlife Service listed it as endangered under the federal Endangered Species Act. As the curator of the University of California Botanical Garden at Berkeley, I work with the national Center for Plant Conservation and a coalition called California Plant Rescue. Each year we make an ambitious plan for conservation fieldwork in the greater San Francisco Bay Area, and for 2020, we planned a packed calendar. Most of our fieldwork was derailed by the restrictions put in place to limit the spread of COVID-19, especially given the timing of the restrictions. Annuals and herbaceous perennials on California's Central Coast tend to have a short spring cycle of growth and seed set. By the time permission was given to be in the field for just day trips, seeds had already set and been dispersed for many species. Scotts Valley polygonum, in contrast, is an annual wildflower that typically starts to germinate in December, flower from May to August, and set seeds in August. The species is now known to occur on less than an acre of A Conservation SOS: Polygonum hickmanii Holly Forbes DAVID GREENBERGER In recent years, the endangered Scotts Valley polygonum (above) has been observed in only one wild population. FORBES, H. 2021. A CONSERVATION SOS: POLYGONUM HICKMANII. ARNOLDIA, 78(3): 5-6 6 Arnoldia 78\/3 \u2022 February 2021 private land adjacent to a new housing development. The development company established a conservation easement to protect Scotts Valley polygonum and another endangered species, Scotts Valley spineflower (Chorizanthe robusta var. hartwegii). Both species are in the buckwheat family (Polygonaceae). In 2015, no Scotts Valley polygonum were found at this site, and it wasn't until 2020 that the number of plants went above four hundred, less than 25 percent of the population observed in 1997. In the past, Scotts Valley polygonum has been documented at two nearby locations, but no specimens have been observed there in recent years. One of these locations is a special ecological preserve adjacent to Scotts Valley High School, where the polygonum has not been observed since 2015. The site is fenced and managed to support the species, but we have limited hope it will reappear on its own. When my colleagues and I could finally return to the field, pandemic protocols required all participants to travel solo in vehicles and to maintain at least a six-foot distance from one another when working at the sites. I was fortunate to work with two other botanists, Kathy Lyons and Jaymee Marty, at the easement site on August 7. We declared ourselves free of COVID-19 symptoms and signed liability waivers for the landowner. The plants occupied an area of less than forty square feet, scattered across an undulating grassland. We worked for hours on hands and knees making a modest seed collection from the less than five hundred plants\u2014all that is left in the world. As we collected the tiny seeds from the plants (removing only a small percentage of the seed set), we remarked on how it almost felt normal to be in the field again, despite the pandemic. Travel restrictions had resulted in a huge reduction in the number of cars on the road, which meant that, as a side benefit, travel between Berkeley and Scotts Valley flowed along at the speed limit, instead of crawling through typical Silicon Valley gridlock. Travel each way took one hour instead of the usual three. A few weeks after our work, the CZU Lightning Complex wildfire in Santa Cruz and San Mateo Counties blackened over eighty-six thousand acres, starting on August 16 and continuing through September 22. The evacuation zone included the two historic polygonum sites. The only extant site, from which the seeds had been collected, was on the margin of the evacuation zone, just across a four-lane highway. It could have easily been different. The fire burned so hot in places that any seeds present in the soil were cooked. During the fire we anxiously checked the maps. It was a great relief to learn that the polygonum sites did not burn. Our purpose for collecting seeds was twofold: first, to create a conservation seed bank as a backup in case the population is lost for any reason, and second, to produce more seeds by growing plants in a nursery environment. This amplification of seed numbers may make it possible both to reestablish the plants at their historic sites and to augment the numbers of plants within the conservation easement. In November, propagator Susan Malisch at the University of California Botanical Garden sowed one-third of the polygonum seeds from our seedbank. As of late January 2021, over 85 percent germination has been observed. Each seed was sown individually to minimize root disturbance when the plants are moved into larger containers. The plants aren't likely to grow larger than six inches tall and perhaps two inches across\u2014giants compared to the plants in habitat, where they are crowded together and typically grow about one and a half inches tall. We look forward to a successful crop of Scotts Valley polygonum in 2021. If all goes as planned, we will have thousands of seeds to use in saving this species from extinction. Wildfires and other threats still pose an incredible risk to the species, but with a robust conservation seedbank and the knowledge of how to grow the plants to reproductive size, we can safeguard its future. Botanists are paying close attention, and Scotts Valley polygonum is no longer overlooked. Next November, we plan to work with the federal Recovery Implementation Team\u2014 a team established by the Fish and Wildlife Service\u2014to place seeds back into the habitat. Holly Forbes is the curator of the University of California Botanical Garden at Berkeley. Support for the Scotts Valley polygonum project is provided by the Ventura Office of the United States Fish and Wildlife Service."},{"has_event_date":0,"type":"arnoldia","title":"An Unusual Autumn at the Dana Greenhouses","article_sequence":3,"start_page":7,"end_page":9,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25722","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24e8927.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Enzenbacher, Tiffany","article_content":"October was quiet. The headhouse at the Dana Greenhouses was still, except for the dim hum of the radio, a necessity for an almost empty building. In previous years, the same location would have been marked with a cacophony of sounds, the door thrown ajar as Arnold Arboretum plant collectors eagerly arrived to unpack their hard-earned seeds and plants. Sieves and colanders would have rattled against the center worktable as plant production staff removed fruit pulp from each seed, and everyone would be talking about new and exciting acquisitions. Seed cataloging and cleaning is a departmental undertaking, sometimes lasting the entirety of fall and into early winter. This annual activity has occurred at an invigorated level since 2015, when the Arboretum launched the Campaign for the Living Collections, a strategic ten-year initiative to increase the biodiversity and conservation holdings of our living collections by adding nearly four hundred wild-collected taxa that were not already growing in our landscape. As part of the campaign, staff organized and executed as many as five expeditions annually, traveling to locations in northern Idaho, central China, the country of Georgia, and elsewhere. I have participated in two of those expeditions myself: one to the Ozarks and another to northern Illinois and Wisconsin. It was rewarding to engage in the full process, from planning An Unusual Autumn at the Dana Greenhouses Tiffany Enzenbacher The pandemic changed fall and winter routines at the Dana Greenhouses, providing an unplanned reprieve from processing new, wild-collected plant material. Chris Copeland (above) prepares grafts of a plum (Prunus alleghaniensis), one of hundreds of clonal propagations that are completed annually. ENZENBACHER, T. 2021. AN UNUSUAL AUTUMN AT THE DANA GREENHOUSES. ARNOLDIA, 78(3): 7-9 TIFFANY ENZENBACHER 8 Arnoldia 78\/3 \u2022 February 2021 expedition logistics and obtaining permits to harvesting in the field and then processing seed back at the Dana Greenhouses. The collection that stands out most from my two experiences was of the endangered seaside alder (Alnus maritima ssp. oklahomensis). I collaborated with Kea Woodruff, then the Arboretum's plant growth facilities manager, to collect seed from two plants growing along the Blue River in Tishomingo, Oklahoma. We were guided by local experts. This subspecies of the seaside alder has only been documented in three other locations in the wild, all near the Blue River. (The two other subspecies also have extremely restricted ranges\u2014one occurs in a single location in northwestern Georgia, the other com- When new plant material arrives at the Dana Greenhouses, staff begin a detailed process of record keeping. New innovations have streamlined the process. Sean Halloran (above) readies softwood cuttings and will note rooting observations using a newly developed mobile application in spring. prises scattered populations on the Delmarva Peninsula of Delaware and Maryland.) For me, this collection brought home the purpose of the campaign and the urgency of preserving threatened taxa. In the fall of 2020, however, those collections ceased due to the pandemic. Planned expeditions to China, Japan, and South Korea were postponed. In the headhouse of the Dana Greenhouses, the difference was striking. Only two or three members of the plant production department worked on-site on any given weekday, in an effort to de-densify our workspace and to allow staff to care for children who were completing schoolwork from home. This revised schedule continues into the new year. Other TIFFANY ENZENBACHER Dana Greenhouses 9 nonessential staff are not permitted inside the building. Now, our team hears only the quiet sounds of greenhouse doors opening as we check the facilities, monitor plants for water, and scout for insect pests and diseases. We hear the clatter of containers being placed on potting benches as we prepare to transplant seedlings and the swish of cutting media components being mixed as we get ready for winter hardwood cutting season. We occasionally share the same workspace, but only brief, work-related interactions can take place. Our team meetings are now virtual. The production cycle for plants already in the greenhouses and nurseries has not significantly slowed this year, although the headhouse tables are bare: no collection sheets from the expeditions strewn about, no bags of fermenting berries or cones to go through. During this altered time, as we have continued with usual greenhouse and nursery tasks, the plant production department has had the opportunity to refocus our direction on other activities. We have made enormous strides to integrate our workflows into the Landscape Management System, a new digital tool developed at the Arboretum, which combines horticulture and curation efforts through mobile applications and an internal website. One component of this system, PropManager, will eventually replace the use of handwritten propagation cards, which are used to record treatments and results for propagation attempts, including for seeds that return from expeditions. Currently, when seeds arrive, staff record propagation methods and experiments on these cards. While some seeds can be sown immediately, others must undergo periods of cold or warmth. Others require treatments to weaken the seedcoat: sandpaper or an acidic solution. Data from propagation cards are then entered into BG-BASE, the Arboretum's plant records database. Then, as germination, transplanting, and other events occur, the cards are updated, corresponding data are input into BGBASE, and the cards are refiled into a binder. PropManager will allow us to create a digital \"card\" on a mobile device and record events in real time. We observed how inefficient the physical card system was when Sean Halloran, our plant propagator, had to transport boxes of binders to and from his home as he toggled between remote and on-site work this spring. Our team has also completed work that will help us to map, track, and communicate about plants in our nurseries using additional Landscape Management System tools. Chris Copeland, our greenhouse horticulturist, worked with members of the Landscape Management System team to acquire and upload locations of over 250 nursery plants. Specimens are now visible on a dynamic map, and we can easily picture spatial patterns and adjust maintenance of the next generation of Arboretum plants. Likewise, when horticulture staff inherit a tree after it has been transplanted into the landscape, they can use this new set of tools to determine noteworthy events that transpired during the tree's early life. We are also working with Mike O'Neal, the director of BG-BASE, to analyze information about our repropagation attempts. Each year we duplicate hundreds of historic Arboretum plants through vegetative propagation\u2014a process whereby resulting progeny are genetically identical to the original. Halloran and O'Neal are in the process of creating BG-BASE summary reports. The result will help determine whether the repropagation of a specimen in the landscape is complete. Instead of Halloran spending weeks at his desk writing code and manually sleuthing through BG-BASE tables, he will be able to run a quick query to have access to all the data needed. The scene at the Dana Greenhouse is certainly different than it was in autumn 2019. That year, we processed over 150 seedlots and mailed surplus material to over a dozen collaborating institutions. Yet the unplanned reprieve from receiving campaign material has allowed our plant production team to collaborate on projects that would have otherwise progressed incrementally over multiple years. We are now better equipped than ever and prepared for the onslaught of new seed collected by Arboretum explorers who are eager to be back out in the field. Tiffany Enzenbacher is manager of plant production at the Arnold Arboretum."},{"has_event_date":0,"type":"arnoldia","title":"A Brief History of Juglandaceae","article_sequence":4,"start_page":10,"end_page":17,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25723","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24e896b.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Frei, Jonas","article_content":"When I first encountered butternuts on the ground of the arboretum here in Z\u00fcrich, Switzerland, I was puzzled. The tree these nuts fell from must have died or been felled years ago, so I only had the seeds for identification. This North American species, Juglans cinerea, is rarely seen in European cultivation outside specialized tree collections, and I didn't recognize the ridged, oblong nuts. When I took a few home, they were not easy to identify within books on common park trees. After additional research, however, the butternut aroused my fascination and left me with questions about the whole walnut family (Juglandaceae). I had long been familiar with this group of plants, but the more I read about them, the more I realized that, in fact, I knew so little. Like the butternut, many other members of the walnut family were absent in books that I had at home: hickories (Carya), wingnuts (Pterocarya), and platycarya (Platycarya). As I encountered each new species, new questions arose. After several years of intensive study, my pursuit evolved into a book project, Die Walnuss, which was published (in a German edition) in late 2019. My work with this unique plant family went far beyond scientific analysis; it also involved an artistic exploration of the unique variety of forms of this plant family. I wanted to make the knowledge hidden in scientific papers accessible through a language of drawings and photographs. These different approaches\u2014science and art\u2014offered new ways of observing and understanding the world of walnuts. I live in a region with no native species of this widespread plant family. Here, you can occasionally find the North American eastern black walnut (Juglans nigra) planted as an ornamental tree in parks. The English walnut (Juglans regia) was most likely introduced by the Romans into the northern parts of Europe and can often be found growing as lone specimens on farms. But the number of these solitary trees has declined in the region since the industrialization of agriculture half a century ago. Walnut farms and orchards are relatively new in the Germanspeaking part of Europe, and walnuts bought in grocery stores here mostly originate from France (P\u00e9rigord and Grenoble), the United States (California), or Chile. Members of Juglandaceae, however, were once among the most common trees of alluvial forests in Central Europe. Fossils allow us to look back on a plant family whose greatest diversity and distribution preceded the ice ages in the Paleogene and Neogene. Many species disappeared only a few hundred thousand years ago. I became fascinated by this history. The fossil record reveals a long, slow story of evolution and shifting ranges, and it provides a counterpoint to the story of the family's rapid globalization in recent centuries. Not far from Strasbourg, in the Rhine Valley of France, researchers and fossil collectors have discovered fossilized butternuts, described under the name of Juglans bergomensis. These fossils correspond so closely to the North American butternut that it is hard to find visual differences. The nuts must have fallen into the shallow water and sandy substrate of the Rhine five million years ago, but they still have almost the weight and feel of fresh nuts due to carbonization. In fact, this species had a wide distribution: its fossils have been reported in Italy, the Netherlands, and wider parts of eastern Europe and Russia. Similar fossils dating to the Neogene have been found in Japan and in the southern United States. Fossilized hickory nuts are also present in the Rhine sediments, including those of a widespread fossil taxon called Carya globosa, which is similar in appearance to the water hickory (Carya aquatica). Although all the European hickory species went extinct millions of years ago, A Brief History of Juglandaceae Jonas Frei FREI, J. 2021. A BRIEF HISTORY OF JUGLANDACEAE. ARNOLDIA, 78(3): 10-17 Facing page: The walnut family is best known for nut-bearing species like the English walnut (Juglans regia), pictured here in the Thur Valley of Switzerland, but the family also includes notable wind-dispersed species. ALL IMAGES BY THE AUTHOR the nuts look as fresh as if they were only a few years old. Walnut family species with large, animaldispersed fruits are only part of the story. Wingnuts (Pterocarya)\u2014a genus that is now known for six extant species\u2014were once dominant trees here in Central Europe along rivers and in mountain slope forests. These are ancestors of the species we now call the Caucasian wingnut (P. fraxinifolia), which today runs wild in parks and gardens in Central Europe, its root sprouts forming dense stands. Some horticulturists have argued that we should cease planting this species in our gardens, given these invasive tendencies, but based on the fossil record, we could also view the wingnut as a returnee from another era. After all, wingnut leaf fossils in the Stuttgart region were found in sediments of the Holstein interglacial and date back only 325,000 years. The few remaining populations of this once widely distributed species are increasingly threatened in their last refuges in the Caucasus. Wheel wingnuts (Cyclocarya) and platycarya\u2014both unusual wind-dispersed genera now found only in East Asia\u2014are also represented in the fossil records in Europe. The reason the walnut family went extinct in Europe while some species meanwhile survived in North America and East Asia is related to the geographical shape of the continents. Here in Europe, the Alps and the Mediterranean Sea form a barrier for the north-south migration of plant species. In cold periods, trees could survive only in the southernmost corners of Europe; therefore, while in America plant species could migrate according to climate conditions, many European species died out with every cooling and warming. The fossil record indicates that wingnuts survived this back and forth the longest of all Juglandaceae, but in the end, they vanished irretrievably, just like the European magnolias (Magnolia), kiwis (Actinidia), and sweetgum (Liquidambar). Other genera of woody plants, including maples (Acer) and ashes (Fraxinus), are now represented in Europe with only a few species but had much greater diversity before the Pleistocene ice ages that started about two and a half million years ago. The diversity of these genera in Europe was similar to their modern-day representation in North America and Asia. The fossils reveal more than former distributions and long-extinct species\u2014the record also documents how the walnut family evolved from an entirely wind-dispersed family to one with the charismatic nut-bearing species that we know today. Some of the oldest fossils of Juglandaceae fruits originate from the United States. Fruits of a wheel wingnut named Cyclocarya brownii have been found in different sites from the Paleocene, occurring shortly after the K-T boundary, the geologic marker that separated the Cretaceous and Paleogene a good sixty-five million years ago. This event of mass extinction was both the end of the era of dinosaurs and ammonites and the beginning of a new chapter for the walnut family. Cyclocarya looks very typical for early members of the family, especially since its fruits are spread by the wind and not by birds or mam- Fossils document the former abundance of the walnut family in Central Europe, where no members of the family naturally occur today. Hickory (Carya) fossils, shown above, were collected from sediments in the Rhine Valley, close to Strasbourg, France, and are around five million years old. The author's illustrations show both the diversity and beauty of the walnut family: (a) English walnut, Juglans regia; (b) little walnut, J. microcarpa; (c) Japanese wingnut, Pterocarya rhoifolia; (d) Japanese heartnut, J. ailantifolia var. cordiformis; (e) black walnut, J. nigra; (f) butternut, J. cinerea; (g) Arizona walnut, J. major; (h) Platycarya strobilacea; (i) Ma walnut, J. hopeiensis; (j) Manchurian walnut, J. mandshurica; (k) nutmeg hickory, Carya myristiciformis; (l) buart hybrid, J. x bixbyi; (m) Chinese butternut, J. cathayensis; (n) bitternut, C. cordiformis; and (o) Chinese wingnut, Pterocarya stenoptera. mals. Back in Paleocene, some fifty million years ago, mammals only started to specialize in the new ecological niches that became available after the extinction of the dinosaurs. Many other winged walnut species emerged. Some went extinct, but the descendants of others are now populating the tropics of the New and Old World: Oreomunnea in Central and South America, and Engelhardia in Southeast Asia and northern India. It was only with the diversification of mammals, especially squirrels, that some walnut species developed fruits that could be spread by animals. Squirrels and other rodents drove the evolution of Juglandaceae in two different genera: walnuts (Juglans) and hickories (Carya), which evolved within separate lineages. Birds, especially the crow family, likely played a part in the distribution from the beginning as well. Because animals never find all the nuts they stash in their winter storage places, they contributed to the spread of these groups, and evidently, they were quite efficient. Walnuts and hickories spread through North America, Asia, and Europe, populating much of the Northern Hemisphere. In the case of the walnuts, this process must have taken place during the span of about ten million years. The oldest known fossil record of the genus, a species named Juglans clarnensis, was discovered in North America and dates back forty-four million years, while the oldest European specimen of J. bergomensis is around thirty-three million years old. Later, humans helped with the worldwide spread of two major species: the English walnut and the pecan (Carya illinoinensis). Whereas squirrels and crows spread walnuts and hickories on three continents over several million years, humans extended the range of cultivation into all other suitable climatic regions within a few decades. The English walnut (a species of Eurasian origin) and pecan (from the southeastern United States) are now cultivated well outside their native range, including in parts of South America, northern and southern Africa, Australia, and New Zealand. So, the tasty kernels of the walnut became the main reason for this widespread distribution\u2014a process started by squirrels many millions of years before the fossil records prove the evolution of humans. Today, in Central Europe, almost forty species and hybrids of Juglandaceae are cultivated. During my research, I traveled to many parks and arboreta, looking for insight into the diversity of this family. I was driven not only by my scientific interest in Juglandaceae but also by my enthusiasm for the aesthetics of their habits, leaves, and fruits. The readers of my book should be able to make their own journey of discovery through the walnut family, on the tracks I have uncovered with my research. Often, after days of traveling, I would find out that a tree I wanted to visit had been cut down or that a rare species was simply confused with an ordinary, oft-planted one. I created a collection of seeds of all the cultivated species and a leaf herbarium. The collection soon included hundreds of fruits and nuts from different locations in Europe, which made it possible to distinguish between the species and hybrids. Later, the collection became the basis for the illustrations of all species in the individual portraits of the book. These trips through Europe searching for the different species of the walnut family also brought to light the stories of other humans\u2014 botanists and horticulturists\u2014who moved the walnut family all over the world. While I could find many species within a day or two of searching, many researchers spent years traveling through the natural habitats in North America and Asia a few centuries ago. In the time of Carl Linnaeus, only three walnut species were known to European researchers. Besides the English walnut, Linnaeus included the North American butternut and the eastern black walnut in his Species Plantarum, published in 1753. The hickories\u2014especially the Asian species\u2014were documented much later. The genus name Carya was proposed by the English botanist and plant collector Thomas Nuttall, who used the name, in 1818, in his work The Genera of North American Plants. He had borrowed this name from ancient Greek, where karya was a word for walnut. The valid botanical name for a genus or species should always be the one from the first official description, and in this case, Nuttall's proposal wrongly became the namesake of the genus. Ten years earlier, the hickories were described under the name Hicoria by the American polymath Constantine Rafinesque. These circumstances led various scientists to urge for reinstating the earlier name, but the change was never implemented. It would have been a respectful act, not only to honor the scientific rules but also because the Greek word karya refers to the English walnut whereas Hicoria is derived from the Algonquin word for a well-known hickory dish: pocohiquara. That name reveals an obvious fact: these trees have a cultural importance that far predates their scientific documentation. Philipp Franz von Siebold was one of the first Europeans to collect plants in Japan. One of his great collections was Platycarya strobila- The large kernels of walnuts and hickories have inspired animals to disperse the species widely: (a) English walnut (cultivar), Juglans regia; (b) water hickory, Carya aquatica; (c) shellbark, C. laciniosa; (d) Chinese hickory, C. catha- yensis; (e) bitternut, C. cordiformis; (f, g, h) English walnut (cultivars); (i) butternut, J. cinerea; (j) black walnut, J. nigra; (k) Japanese walnut, J. ailantifolia; (l) Japanese heartnut, J. ailantifolia var. cordiformis; and (m) pecan, C. illinoinensis. cea, which was described in 1843. Some botanists initially thought it was a conifer due to its cone-like fruiting structures. In 1844, the famous English plant collector Robert Fortune also found Platycarya in China. Assuming that it was a new, not-yet-described species, he sent herbarium material and seeds to the Royal Horticultural Society in London. John Lindley, the secretary of the society, named the plant after its finder, Fortunaea chinensis, and called the species the most important new find of Fortune. Later, it became known that Siebold had described the species one year earlier, so today the name Fortunaea is only used as a synonym. These scientific explorations\u2014and those of other botanists\u2014made it possible to describe, collect, and, of course, cultivate many of the species as ornamentals and orchard trees. But this era of Siebold and Fortune was not simply a time of great scientific discovery; it was also a time of European colonization, in which the gathering of knowledge on expeditions was often combined with ideological, cultural, and religious imperialism. This movement of plants around the world coincided with violations of ethical standards by European maritime powers and a merciless approach to other cultures. The relatively slow but efficient distribution of Juglandaceae by squirrels and mice seems innocent in comparison. When the walnut family is viewed in the broad sweep of its evolutionary history, the speed of its recent spread is clearly unprecedented. As beautiful as it is to see the worldwide diversity of Juglandaceae close together in many parks today, the globalization of the family has also produced novel threats. As humans moved the walnut family around the world, fungi and pathogens often migrated with the species. In the United States, a fungal disease known as the butternut canker (Sirococcus clavigignenti-juglandacearum) has brought the butternut to the brink of disappearance. The fungus, which was once native to Asian walnut species, causes little damage to its original hosts, but it is often fatal to the North American butternut. The thousand cankers disease, meanwhile, is the result of the unfortunate encounter of a fungus (Geosmithia morbida) and a beetle (Pityophthorus juglandis) that formed in the western United States due to the proximity of the eastern black walnuts, cultivated in parks, and natural populations of the Arizona walnut (Juglans major). And the walnut fruit fly (Rhagoletis completa), which once lived inconspicuously on the black walnut species of North America, today spreads quickly in walnut orchards of Europe. Meanwhile, the close planting of related Juglandaceae species leads to the formation of hybrid offspring. This has led to major changes in natural environments, especially in the case of the butternut populations in North America. Many of the butternut trees that can resist the butternut canker also carry the genetic material of Japanese walnuts (Juglans ailantifolia). Resistant hybrids have greater fitness, as they survive and have more offspring, which could be a blessing for the American butternut stocks that survive the strong fungal infestation. On the other hand, conservation of the \"real\" butternut becomes more complicated. This scenario reveals the cascade of unintended but profound environmental consequences of human actions, which cannot be easily resolved. Of course, the walnut family experienced various climatic changes over the past fifty million years and therefore changed its distribution again and again. It is assumed that many of the species we know today are the result of hybridization between different populations that collided after a long separation due to climatic fluctuations and subsequent spread by squirrels and ravens. Genetic studies suggest that the English walnut originated from the hybridization of the black walnuts (section Rhysocaryon) and Asian butternuts (section Cardiocaryon). Also, the American butternut is said to carry some black walnut genes in addition to the genetic material of similar Asian species from the Cardiocaryon section. Given this history, one could say that many walnuts, as a lineage, will adapt to human-made influences, although it is unlikely all of the walnut species we know today will survive the pressure. Recently, in a second-hand bookstore, I found a small booklet titled Die Quaianlagen von Z\u00fcrich, from 1889. The author, botanist Carl Joseph Schr\u00f6ter, planned the tree collection at the arboretum where I first encountered the butternuts that started my interest in this exceptional plant family. He states that a butternut tree was planted in 1887 at exactly the spot where I found the nuts pressed into the soil. Now I know that these nuts, almost like modern-day fossils, are the remains of a nowrare species. The tree was planted long before butternut canker was imported to the United States, and before hybridization with imported species changed its natural populations rapidly. If we did not have our own hands in all the processes that threaten species like the butternut, we could analyze the consequences from a scientific perspective and see with great fascination how some species emerge from this immense pressure and how others disappear, just like during the whole history of this family. But we also have a responsibility towards biodiversity, towards those species that exist now and that enriched the global ecosystem long before the arrival of humans. Today, as the pace of ecological change and movement continues to accelerate, we have to recognize that the story of the walnut family is now entwined with our own. Jonas D. Frei is a landscape architect, documentary filmmaker, illustrator, and author from Z\u00fcrich, Switzerland. The author's book Die Walnuss\u2014currently available in a German-language edition\u2014features drawings, photographs, and descriptions of the walnut family."},{"has_event_date":0,"type":"arnoldia","title":"Discovering the Majestic Mai Hing Sam of Laos","article_sequence":5,"start_page":18,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25724","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25ea36e.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Coffman, Gretchen C.","article_content":"Facing page: The author was the first researcher to document the critically endangered Asian swamp cypress, Glyptostro- bus pensilis, growing in Laos. This old-growth specimen, photographed in 2015, is locally known as the \"mother tree.\" PHOTO BY DAVID MCGUIRE In early April 2007, less than three weeks after submitting my dissertation and receiving my doctorate at the University of California, Los Angeles, I got on a plane headed for Laos. It took four flights and more than twenty hours of flying time to get to the capital city of Vientiane. From there, I was bound for the Annamite Mountains: an eight-hour drive from Vientiane, then onward by truck, hand tractor (tok tok), and boat. The remote Annamite Mountains run 680 miles (1,100 kilometers) along the border between Vietnam and Laos, reaching into northern Cambodia. This range divides the Mekong River Basin to the west from Vietnam's narrow coastal plain to the east. The mountains are home to exceptional biodiversity. After the Vietnam War ended, Laos closed to Westerners, but in the early 1990s, the borders began to open. Biologists began to document fascinating endemic wildlife, some new to science, including the enigmatic saola (Pseudoryx nghetinhensis), a critically endangered bovine that, due to its rarity, has been dubbed the Asian unicorn. Perhaps the most miraculous discovery was that of the endangered Laos rock rat (Laonastes aenigmamus), a rodent identified as a surviving member of a family (Diatomyidae) previously thought to have gone extinct about eleven million years ago.1 Plant biodiversity in this mountain range is exceptionally rich as well, and many new species have been documented.2 When I initially arrived in the foothills, I could not have imagined that I would become part of one of these discoveries: the first biologist to collect samples of the majestic Asian swamp cypress (Glyptostrobus pensilis) growing in the country. This critically endangered species\u2014locally known as mai hing sam\u2014is currently documented in only two other heavily degraded populations, both in Vietnam. The mai hing sam in Laos are the only old-growth specimens in the world, and in recent years, the stands have been increasingly threatened by agricultural development and poaching for the luxury timber market. The protection of the few hundred remaining individuals in Laos has become my mission. Arriving in Laos My journey to the Annamite Mountains had begun four months earlier, when a member of my doctoral committee, Phil Rundel, emailed me with a proposal to work on a project in an especially remote part of Laos. I was immediately intrigued by the biodiversity, and the thought of getting away from my computer days after finishing my dissertation was alluring. Yet, I was hesitant. The opportunity involved working as a restoration ecologist on a World Bank hydropower project. As a wetland and riparian ecologist by training, I had always focused my research and professional work on protecting rivers and streams, not damming them. Rundel encouraged me to research both points of view\u2014pro- and anti-hydropower dam. On my breaks from dissertation writing that winter, I read articles and websites from advocates and opponents (including, among the latter, International Rivers and other nongovernmental organizations). I also corresponded with wildlife biologists who would be working on the project. The work was part of mitigation actions for the Nam Theun 2 Hydropower Project and supported the development of a national park in the reservoir's headwaters. At more than 1,300 square miles (3,500 square kilometers), this protected area is one of the largest remaining contiguous areas of forests on the Indochinese Peninsula.3 Ultimately, I made a pragmatic decision: there was no stopping the dam, but I could work for the wildlife by helping to develop a conservation plan. I would work closely with 20 Arnoldia 78\/3 \u2022 February 2021 The Annamite Mountains\u2014known for complex topography, geography, and climate\u2014harbor some of the most-contiguous moist forests in Indochina. James Maxwell, a renowned botanist from Chiang Mai University in Thailand, along with a team of wildlife biologists from a multitude of disciplines. Our mission was to assess wetland habitat on the Nakai Plateau\u2014located high within the Annamite Mountains\u2014before it was flooded by the reservoir. We would document the wetland vegetation and develop a wildlife management plan that included the restoration of habitat within an area known as the Nakai- Nam Theun National Protected Area. Little did I know I would be acting as field coordinator once I arrived, a task that I was comfortable with from fifteen years of managing restoration projects in the United States but not nearly as easy in this new landscape and culture. The Discovery The Annamite Mountains contain some of the last relatively intact moist forests in Indochina, unique due to the region's complex geology and climate, and relatively inaccessible due to the steep topography. Initially, working with Maxwell proved extremely difficult. He could not understand why I had been hired on this project, since all my botanical experience was in the United States. He was standoffish and focused on collecting rare wildflowers he encountered. As we settled into the work, however, we bonded. He proved to be an exceptional mentor and friend, and in the years to come, I would stay with Maxwell and his wife in Thailand on multiple occasions. Our standard workdays were reminiscent of my first fieldwork experiences in the hot, humid wetlands of coastal Georgia, where I had grown up. When we arrived in Laos, it was the height of the dry season and unbearably hot in the late afternoons. We started at sunrise to avoid the heat, first eating a bowl of pho, a noodle soup loaded with fragrant mint, crunchy cabbage, long beans, and assorted leathery forest leaves. In the field, we lugged our plant presses everywhere, as everything we collected ASSOCIATION ANOULAK Glyptostrobus 21 The author located Glyptostrobus pensilis within the Nakai-Nam Theun National Park. The discovery was made while assessing wetland habitat and developing a wildlife management plan for the Nam Theun 2 Hydropower Project. ARNOLD ARBORETUM, ESRI, GARMIN, AND GIS COMMUNITY went immediately into the press. The afternoons were sticky and oppressive in the open wetlands. We ended around four o'clock when we couldn't take the heat anymore, giving us time to process our plant specimen and clean up our notes. At that point, the plants went directly from the presses into rice sacks with alcohol for preservation. We surveyed all the herbaceous wetlands across the Nakai Plateau. These wetlands intermingled with rice paddies and were often used as grazing pasture. We began our collections in large, easy to access wetlands on the south side of the Nam Theun River. To guide us, we used paper topographic maps. We then made our way to more forested wetlands and riparian forests, northwest towards the dam site and onward to an area that was nicknamed Thousand Islands because of how the landscape fl ooded during the monsoon rains. From there we continued east, across the river, near the foothills of the Annamite Mountains. The fi rst potential wildlife habitat restoration site we visited was northeast of Thousand Islands, near the Nam Xot tributary to the Nam Theun River. Our colleague Pierre Dubeau, a geospatial scientist who had sited these potential restoration areas, exuberantly walked downstream through the forested wetland toward an area with large wetland grasses (Neyraudia reynaudiana). Maxwell and I followed Dubeau and wildlife biologist Rob Timmins, who was carrying an umbrella in the sprinkling warm afternoon rain. We agreed that this would be a great open location, ideal for wildlife habitat restoration. As we trudged back among a mucky mess of the forested wetland swamp, I stumbled over something and fell to my hands in the soggy soils. I slowly got up, shook off the fall, and investigated what I tripped over. It looked like a pneumatophore\u2014the cypress knees I knew from my childhood in coastal Georgia, where bald cypress (Taxodium distichum) are a dominant feature of the swamps. SOUTH CHINA SEA Mekong River VIETNAM THAILAND LAOS Route 13 A N N A M I T E M O U N TA I N S Nakai-Nam Useun National Park Phou Hin Poun National Biodiversity Conservation Area kongngngngngngn R 25 miles 50 kilometers Nam Useun Reservoir Protected Areas Glyptostrobus 23 Facing page: Forests in the Annamite Mountains are rapidly disappearing due to forestry, agriculture, and hydropower development, along with other causes. Philip Thomas (right) stands beside Glyptostrobus pensilis within a rice paddy. PHOTO BY DAVID MCGUIRE I looked up to find the tree it might be attached to, and sure enough, an enormous conifer towered above me. I looked up at this red-barked giant and saw something wonderfully strange and familiar. It looked like a cross between the bald cypresses that I knew from Georgia and the coastal redwoods (Sequoia sempervirens) from California, both members of the cypress family (Cupressaceae). I found several other knees as I walked up to inspect the tree. This, I proclaimed to Maxwell, must be a very special tree! Maxwell, however, like many other tropical botanists, was not as interested in conifers as much as the epiphytes that might grow on them. He thought nothing of it. Meanwhile, I collected the samples of small cones, foliage, and bark of this tree, which I sent to conifer expert Philip Thomas at the Royal Botanic Gardens, Edinburgh, for identification. Documenting the Mai Hing Sam Conifers are dominant or codominant parts of primary- and secondary-growth evergreen forests throughout the Annamite Mountains. In Vietnam, for instance, the mountains host a particularly rich assemblage of thirty-three conifer species, of which the cypress family (Cupressaceae) has seven.4 When I asked people in the neighboring Lao communities about the enormous tree that I had encountered, they provided a name: mai hing sam. Mai means \"tree,\" hing is a modifier for the kind of tree, and sam means \"swamp,\" or what ecologists would describe as a forested wetland. As it turned out, the mai hing sam was, indeed, special. When Philip Thomas replied to my email, he identified the species as Glyptostrobus pensilis (known as the Asian swamp cypress), which the International Union for Conservation of Nature has classified as critically endangered.5 In 2007, the scientific community was aware of only 250 individuals of this species in the wild in Vietnam, where most were spindly, unhealthy young trees, growing in two small stands in the middle of coffee and corn plantations. Other stands in China were presumably planted.6 Due to its rot-resistant wood, Glyptostrobus pensilis is highly sought after in the luxury timber market and is used for a variety of structural and boat-building uses by local communities. It is threatened (like so many endangered species) by illegal logging. As I learned more about the two populations in Vietnam, I realized how remarkable the mai hing sam in Laos really were. The trees in Vietnam grew very close together and, like those in China, appeared like they could have been planted. Boardwalks had been built within the stands to get around. Dams located beneath each of the stands were used for agricultural irrigation and raised the water levels for the trees significantly. In contrast, the trees that we observed in Laos were erect and widely spaced, as expected for a wild population. The crowns of the mai hing sam in Laos were only found in the top third of the trees, with no limbs below for us to climb to the seed-bearing cones. In the Vietnam population, perennial and annual branchlets were numerous along the main bole, appearing to be epicormic growth. This form suggests that the trees in Vietnam were responding to stress from inundation. Also, some of the trees in Vietnam were cut down years ago and had resprouted.7 I immediately told my colleagues about the mai hing sam discovery so that we could develop a strategy to describe and protect this stand. I also informed the Nam Theun 2 Power Company (NTPC) of the discovery and asked to spend time describing the tree and its ecology and to have a surveyor document their elevation relative to the proposed reservoir footprint. I was not allowed time to document this stand properly, however, and I was only able to record the number and size of the trees and basic soil characteristics. There were approximately one hundred trees in the stand, and many were three feet in diameter at breast height. We only had very rough elevation information from our GPS units, but it was clear that the trees\u2014along with many others that we were unable to document\u2014would likely be within the reservoir footprint. In desperation to protect these rare trees, I contacted the Nam Theun 2 Panel of Experts, an audit group that was in charge of assessing 24 Arnoldia 78\/3 \u2022 February 2021 the environmental and socioeconomic impacts of the dam, during their visit to the Nakai Plateau in August 2007. One of the members, the American conservation biologist Lee Talbot, joined me on a tour of this newly discovered mai hing sam stand. Nothing seemed to come of the visit, however, and unfortunately, I didn't find anything about the trees in the panel's next report.8 I proposed to my contacts at NTPC to collect as many seeds as possible and try to propagate and grow more trees. NTPC thought it was a great idea and gave us the go-ahead. Developing a Restoration Protocol At the time, mai hing sam had never been successfully propagated from wild-collected seed. As a result, several critical facts about restoration protocol were unknown to scientists: What time of the year do the seeds mature in the mountains of Laos? How long is their seed viable? Do they produce seeds every year? Did we need to treat the seeds before sowing them? Under what conditions would they propagate and survive? What we did know was that all conifer seeds are wind dispersed, so we hypothesized that their dispersal is probably connected to the windy part of the year, which occurs toward the end of the monsoon season. Our first challenge was logistical: how would we collect seeds from cones high in the canopies, sometimes one hundred or more feet high. Maxwell\u2014who, by this point, had returned to Thailand where he lived\u2014often hired local tree climbers to make collections. But this method requires low branches or woody vines growing up the trunk, as the climbers do not use any specialized equipment. We put our heads together and came up with an unusual plan. We placed large tarps under the trees and hired boys with slingshots to shoot rocks up into the canopies of the trees so that the seeds would fall onto the tarps. We tried this method, and miraculously it worked. We got thousands of cones and hundreds of thousands of minute winged seeds. The next challenge was to clean and propagate the seeds. This process was not managed by a conifer expert like Philip Thomas, as I had hoped. Rather, NTPC hired a commercial contractor to propagate the seeds in a local nursery. The contractor had no familiarity with this sensitive species, and only twelve seedlings germinated. Of those, only four grew to maturity. In restoration and horticultural propagation, this rate is not considered successful, but it was a start. In 2008, NTPC planted the four trees at the confluence of two small streams behind the house occupied by the director of the Watershed Management and Protection Authority. This area was somewhat protected and easy to monitor, although soil characteristics were not similar to the natural conditions of the peat swamps in which the trees naturally grew. In 2015, when I first observed these trees, they were about six feet in height, and on my last expedition, in January 2020, they had reached over sixteen feet. The key to the survival of these four trees, I believe, was sustained high soil moisture during their establishment period and protection using sturdy exclusion fencing to fend off the cattle and water buffalo that munch on the succulent foliage. Threats to Wetland Habitat and Endangered Species After my contract was completed in 2009, I returned to California, where I became an assistant professor at the University of San Francisco. I vowed to go back to look for more mai hing sam in the Nakai-Nam Theun National Protected Area. Southeast Asia is experiencing rapid habitat loss, biodiversity declines, and risk of species extinction primarily due to unsustainable harvesting of forest resources and conversion for agriculture. Lack of enforcement and pressure to develop rice paddies has led to the decline of wetland habitat and continued poaching in the protected areas.9 Nearly every species of softshell turtle, terrapin, or tortoise is threatened with extinction. Populations of exceptionally rare species, such as the saola, are too low and fragmented to be viable.10 Considering these threats, I knew that we needed to mount a concerted effort to document and conserve mai hing sam in the region. Phil Rundel, who had first encouraged me to participate in the project in Laos, recommended that I apply for National Geographic funding. I spent two years getting collaborators on board Glyptostrobus 25 In 2015, the author partnered with other researchers and local collaborators to locate more than six hundred previously undocumented Glyptostrobus in the Nakai-Nam Theun National Park. The author (at right) measures tree height using a clinometer, and a tree climber ascends to the upper canopy. and finding out from contacts if there were any other trees in the national protected area. Maxwell and I corresponded regularly during this period. Likewise, Philip Thomas was a huge source of support and encouragement. Finally, in the spring of 2014, my collaborators and I received funding, and we went on to get permits and work on the expedition plan that summer. With the help of National Geographic funding, we were able to document more than six hundred other mai hing sam between ten and thirty miles from the original stand. These plants occurred in the newly renamed Nakai- Nam Theun National Park, an area that has been under the management of the Watershed Management and Protection Authority since 2005. The trees in the oldest stand are more than three feet in diameter at chest level and five hundred to more than one thousand years old. Many of them are over six feet in diameter, and the largest is over ten feet. (We recorded 11.2 feet\u20143.4 meters\u2014but it's difficult to get the measuring tape behind all the woody vines and strangler figs on the trunk.) The neighboring communities call the largest tree the \"mother tree.\" It is more than 138 feet (42 meters) in height. We believe it could be two thousand years old, but it is not the tallest tree: that claim goes to one we documented at 184 feet (56 meters) tall. While these trees are protected in the park, illegal activities still occur. Sometime between September 2015 and February 2016, two hundred mai hing sam were logged, leaving the PHOTOS BY DAVID MCGUIRE 26 Arnoldia 78\/3 \u2022 February 2021 GRETCHEN C. COFFMAN total known population at approximately four hundred individual trees. This event was deeply upsetting, especially because, as I later learned, the individuals responsible were aware of the conservation importance. The Laos government took the event seriously and not only arrested the local Lao poachers but aggressively pursued the company in Vietnam that had hired them. Fortunately, the neighboring communities protected the mother tree from the poachers. Another factor that might have contributed to its protection is that the oldest trees are often hollow at the base, much like coast redwoods in California. The younger trees have solid trunks that are more desirable to poachers. This event shifted our project's goals and objectives to focus on community-based restoration program and to identify and protect other unknown stands in the region. Each November, between 2017 and 2020, we collected seeds from the remaining stands. In the first two years, we propagated two thousand seedlings; however, many of these did not survive. We have learned a lot about propagation from these trials, and our team is actively developing improved propagation and planting techniques to restore stands of the mai hing sam in strategic areas of the watershed. We are excited to collaborate with colleagues in Vietnam and China to restore populations there as well. The urgency is clear: after the poaching occurred, the government intervened before the logs were removed from the forest. Some of the fallen trees were more than a thousand years old, and now those trunks remain as warnings on the forest floor. With these threats in mind, our work continues, sustained by the promise of the small seedlings. Endnotes: 1 To learn more about recently documented mammal species in the Annamite Mountains: Dawson, M. R., Marivaux, L., Li, C., Beard, K. C., and Metais, G. 2006. Laonastes and the \"lazarus effect\" in recent mammals. Science, 311(5766): 1456-1458. doi:10.1126\/ science.1124187; MacKinnon, J. 2000. New mammals in the 21st century? Annals of the Missouri Botanical Garden, 87(1): 63-66. doi:10.2307\/2666208 2 Recent botanical discoveries in the Annamite Mountains include many new orchid species. Also, Brendan Buckley, from Columbia University, documented remarkable old-growth specimens of another cypress family species, Fokienia hodginsii, growing in Vietnam's Bidoup Nui Ba National Park. The oldest specimens he found are more than twelve hundred years old, and the tree-ring data have supported Brendan's research on long-term climate change in the region, including primary evidence for the fall of the Angkor civilization. Sano, M., Buckley, B. M. and Sweda, T. 2009. Tree-ring based hydroclimate reconstruction over northern Vietnam from Fokienia hodginsii: eighteenth century mega-drought and tropical Pacific influence. Climate Dynamics, 33: 331-340. doi.org\/10.1007\/s00382-008-0454-y 3 Robichaud, W. G., Marsh, C. W., Southammakoth, S., and Khounthikoummane, S. 2001. Review of the National Protected Area System of Lao PDR. Vientiane, Lao PDR: Lao-Swedish Forestry Programme, Department of Forestry and IUCN; Scudder, T. 2020. A retrospective analysis of Laos's Nam Theun The author (right) plants a Glyptostrobus seedling on National Tree Planting Day in May 2019. More than one hundred Laos government officials participated in the event, including Axay Vongkhamsao, head of the environmental division at NTPC (left); Khamthone Vongphachanh (center); and Thong Eth Phayvanh (second from right), the deputy general director of the Department of Forestry and director of the Watershed Management and Protection Authority. 7 It is interesting to note that pneumatophores of the trees in Vietnam measure about 2 feet (0.6 meters) tall on average, similar to those in Laos; however, the pneumatophores were more abundant in Laos, sometimes numbering dozens per tree and usually much shorter. 8 McDowell, D., Scudder, T., and Talbot, L. M. 2007. Twelfth Report of the International Environmental and Social Panel of Experts for the Nam Theun 2 Hydro Project. Vientiane: Lao People's Democratic Republic. 9 For more on the environmental threats in Southeast Asia: Hughes, A. C. 2017. Mapping priorities for conservation in Southeast Asia. Biological Conservation, 209: 395-405. doi:10.1016\/j. biocon.2017.03.007; Sodhi, N., Posa, M., Lee, T., Bickford, D., Koh, L., and Brook, B. 2010. The state and conservation of Southeast Asian biodiversity. Biodiversity and Conservation, 19(2): 317-328. doi:10.1007\/s10531-009-9607-5; Nooren, H., and Claridge, G. 2001. Wildlife trade in Laos: The end of the game. Gland, Switzerland: IUCN-The World Conservation Union; Appanah, S., Shono, K., and Durst, P. B. 2015. Restoration of forests and degraded lands in Southeast Asia. Unasylva, 66(245): 52-62. 10 For more on conservation of the saola: Tilker, A., Long, B., Gray, T. N. E., Robichaud, W., Van Ngoc, T., Vu Linh, N., Holland, J., Shurter, S., Comizzoli, P., Thomas, P., Ratajszczak, R. and Burton, J. 2017. Saving the saola from extinction. Science (American Association for the Advancement of Science), 357(6357): 1248. doi:10.1126\/science.aap9591 The map in this article was created using Esri, USGS, USFS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, lntermap and the GIS user community. Gretchen C. Coffman is a wetland restoration ecologist and a senior lecturer at the National University of Singapore. She teaches wetland restoration ecology, biogeography, applied ecology, and research methods in physical geography. From 2010 to 2019, she taught field ecology courses in the Environmental Science Department, the Masters of Science in Environmental Management program, and the Environmental Studies program at the University of San Francisco. Dedication I dedicate this manuscript to the late James Maxwell (1945-2015). He was an intrepid botanist, fervent collector, a character like no other, exemplar taxonomy mentor, and trusted friend. Glyptostrobus 27 GRETCHEN C. COFFMAN James Maxwell in 2007 2 Dam. International Journal of Water Resources Development, 36(2-3): 351-370. doi:10.1080\/079006 27.2019.1677456 4 Of the thirty-three conifer species documented in the Annamite Mountains in Vietnam, thirteen are members of the pine family (Pinaceae). The cypress and yellowwood families (Cupressaceae and Podocarpaceae, respectively) include seven species each. The yew family (Taxaceae) has five species, and the plum yews (Cephalotaxaceae) have one. In particular, the Da Lat Plateau in central Vietnam has sixteen species of conifers, representing the highest conifer diversity in Indochina. Ninety percent of these, however, are nationally threatened. Loc, P. K., The, P. V., Long, P. K., Regalado, J., Averyanov, L. V., and Maslin, B. 2017. Native conifers of Vietnam\u2014A review. Pakistan Journal of Botany, 49(5): 2037-2068. 5 While Glyptostrobus has few extant populations, the genus has existed for more than one hundred million years, dating back to at least the middle Cretaceous, and was once quite abundant. Fossils of the genus can be found across all of Asia and North America and as far north as Axel Heiberg Island in the Arctic. See: Greenwood, D. R., and Basinger, J. F. 1994. The paleoecology of high-latitude Eocene swamp forests from Axel Heiberg Island, Canadian High Arctic. Review of Palaeobotany and Palynology, 81(1): 83-97. doi:10.1016\/0034-6667(94)90128-7; Vickulin, S. V., Ma, Q. W., Zhilin, S. G., and Li, C. S. 2003. On cuticular compressions of Glyptostrobus europaeus (Taxodiaceae) from Kaydagul Formation (Lower Miocene) of the Central Kazakhstan. Acta Botanica Sinica, 45(6): 673-680; Jahren, A. H. 2007. The Arctic forest of the middle Eocene. Annual Review Earth Planetary Science, 35: 509-540. 6 To read more about the stands in Vietnam and China, see: Averyanov, L., Phan, K., Nguyen, T., Nguyen, S., Nguyen, T., and Pham, T. 2009. Preliminary observation of native Glyptostrobus pensilis (Taxodiaceae) stands in Vietnam. Taiwania, 54(3): 191-212. doi:10.6165\/ tai.2009.54(3).191; Tang, C. Q., Yang, Y., Momohara, A., Wang, H.-C., Luu, H. T., Li, S., Song, K., Qian, S., LePage, B., Dong, Y.-F., Han, P.-B., Ohsawa, M., Le, B. T., Tran, H. D., Dang, M. T., Peng, M.-C., and Wang, C.-Y. 2019. Forest characteristics and population structure of Glyptostrobus pensilis, a globally endangered relict species of southeastern China. Plant Diversity, 41(4): 237-249. doi.org\/10.1016\/j.pld.2019.06.007; Wu, X., Ruhsam, M., Wen, Y., Thomas, P. I., Worth, J. R., Lin, X., Wang, M., Li, X., Chen, L., Lamxay, V. Le Canh, N., and Coffman, G. C. 2020. The last primary forests of the Tertiary relict Glyptostrobus pensilis contain the highest genetic diversity. Forestry: An International Journal of Forest Research, 93(3): 359- 375. doi:10.1093\/forestry\/cpz063; Li, F. G., and Xia, N. H. 2004. The geographical distribution and cause of threat to Glyptostrobus pensilis (Taxodiaceae). Journal of Tropical and Subtropical Botany, 12(1): 13-20."},{"has_event_date":0,"type":"arnoldia","title":"Backyard Climate Solutions","article_sequence":6,"start_page":28,"end_page":37,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25725","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25ea726.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Faison, Edward K.","article_content":"Carbon dioxide levels in the Earth's atmosphere stand today at 415 parts per million, which is significantly higher than concentrations have reached for at least the past eight hundred thousand years. Throughout this time, levels oscillated between 180 and 280 parts per million, until the mid-nineteenth century, when they began an inexorable rise. By the end of the century, if business as usual continues, carbon dioxide levels could be higher than at any time in the past fifty million years.1 Like many other concerned citizens, I have wondered what one person can possibly do to help stem the rise of carbon dioxide levels, warming temperatures, and accompanying species extinctions that characterize our Earth in the twenty-first century. Carbon is a twopart problem: we must simultaneously reduce combustion emissions and increase the removal of atmospheric carbon dioxide. As an individual, I can take action to reduce emissions (use more efficient LED bulbs, drive a more efficient car less often, use airplanes sparingly), but what about the other side of the equation? I have increasingly come to recognize that, as a landowner, the way I steward the vegetation on my property can make a difference to both sides of this problem. I live in a small, residential neighborhood in an otherwise rural part of Connecticut. My property comprises a one-and-a-half-acre lot, about two-thirds wooded. The other third includes a yard (where the kids can kick a soccer ball), the house, and a gravel driveway that can accommodate several cars. Plants on my property, like those growing anywhere else, remove carbon dioxide from the atmosphere during photosynthesis and store it as carbon molecules in wood, roots, and leaves\u2014a process known as carbon sequestration. Yet it's surprising to learn just how much carbon dioxide is removed by the Earth's natural vegetation: about 30 percent Backyard Climate Solutions Edward K. Faison of all carbon emitted each year globally. With changes in the way we manage vegetation, this percentage could increase dramatically.2 Trees are key. An acre of temperate grassland and an acre of temperate forest store a similar amount of carbon in the soil, but a forest stores as much as seventeen to twenty times more carbon in the vegetation than does a grassland.3 Compare an acre of forest to an acre of lawn, and the carbon storage disparity is far greater. When we replace natural forest with fields, lawn, and other less-natural land covers (like roads, parking lots, and buildings), not only do we release huge amounts of carbon once stored in the trees into the atmosphere but we also sequester significantly less carbon going forward. The Carbon in My Trees I became curious about the role of my property in sequestering carbon and how much of a difference simple management decisions could make towards this end. How much carbon is stored in the trees on my property? To answer this question, I measured the diameter of every tree at least five inches in diameter at breast height and then used carbon estimation (\"allometric\") equations devised by the United States Forest Service and researchers from Harvard Forest to estimate the total biomass in the trees.4 Plant tissue contains about 45 to 50 percent carbon, so dividing total biomass in half is a good approximation of the carbon storage in the plants.5 The results: 226 trees storing 84.3 tons of carbon total, including a forty-inch-diameter black oak (Quercus velutina) and a red oak (Quercus rubra) of nearly the same dimension. These big oaks comprise less than 1 percent of the trees on my lot but store a remarkable 13 percent of the carbon. The big oaks are not idle reservoirs of carbon either. A healthy red oak forty inches in diameter may add two-tenths of an inch to its trunk FAISON, E. K. 2021. BACKYARD CLIMATE SOLUTIONS. ARNOLDIA, 78(3): 28-37 30 Arnoldia 78\/3 \u2022 February 2021 diameter each year\u2014an imperceptible increase to even an observant naturalist\u2014but a layer of carbon equal to adding an entire six-inchdiameter tree.6 The amount of carbon stored in the trees across my property is over 50 percent higher than in an average acre and a half of forest in Connecticut.7 The elevated levels can be attributed to the relatively high density of large trees in my woods, for which I have the past owners to thank. In addition to the two large oaks, seven other trees exceed twenty-seven inches in trunk diameter. A typical acre and a half of forest in Connecticut currently contains only one or two trees of this size.8 Ironically, the forest edge associated with residential properties appears to contribute to large tree growth. Trees within one hundred feet of a forest edge (which many of mine are) grow faster and thus are often larger\u2014and store more carbon\u2014than those in a forest interior because of reduced competition for light and greater leaf area.9 Hence, smaller residential properties can be surprisingly important contributors to carbon sequestration. Natural Climate Solutions As a property owner, I have many different options for how to manage the vegetation growing on my lot to increase the removal of carbon dioxide from the atmosphere and to reduce emissions. These practices are collectively referred to as natural climate solutions.10 By choosing not to convert the forest on my property into lawn or field (a practice known as avoided conversion), I refrain from emitting the carbon stored in those trees into the atmosphere as carbon dioxide: 310 tons of it. (Carbon dioxide emissions can be calculated by multiplying organic carbon\u2014in this case, 84.3 tons\u2014by 3.67). Three-hundred-ten tons of carbon dioxide is equivalent to the annual emissions of sixty-one cars.11 These are not insignificant numbers, and when multiplied across hundreds of thousands of small properties, the potential for avoided emissions is notable. When retaining a forest, I have a range of management decisions that will affect the amount of carbon stored in my woods. At one extreme, I could remove all the adult trees and regenerate a young forest. At the other extreme, I could remove an occasional tree for firewood, a practice that falls within the category of reduced impact forest management, or, by practicing wildlands management, I could remove no trees at all. Not surprisingly, the latter scenarios result in a significantly greater amount of carbon storage in my woods than the former scenario. In fact, any tree removal on a property like mine reduces carbon storage below the potential maximum for that site (although it is also true that if I leave all my trees standing, which I mostly do, and obtain my firewood from another source, I transfer that carbon loss to another property). Hence, reduced impact forest management\u2014retaining more trees, particularly large ones, for more time\u2014can make an important difference in the amount of carbon that is retained in a forest.12 Decisions about tree retention in residential areas often involve mitigating risk to power lines. A few years ago, for instance, the power company asked for my permission to cut three healthy trees on the edge of my previous property: a red oak, white oak (Quercus alba), and pignut hickory (Carya glabra), all with trunk diameters of more than thirty inches. Removing three trees would not have resulted in any forest conversion on my property\u2014indeed, there are young, small trees growing underneath these big ones\u2014but the carbon stored on my property would have been reduced by about eight tons, equivalent to the annual emissions of almost six cars. A large tree thirty inches in diameter also removes about seventy times the quantity of pollutants (including carbon monoxide, ozone, nitrogen dioxide, and particulate matter) as a tree three inches in diameter.13 I decided that the trees were a relatively low risk to the powerlines and would provide more benefits if I allowed them to continue to grow and sequester carbon. Wildlands management, the decision not to cut or mow any trees, has obvious limitations near houses, but it can be applied to more removed areas. In the relatively small number of wilderness areas and strict nature preserves in the northeastern United States, the trees store a disproportionately large amount of carbon Backyard 31 relative to the region's total forest area.14 Wildlands also have the potential to sequester much additional carbon. Because of a lengthy land-use history of forest clearance and intensive logging, northeastern forests are, on average, only about 20 to 30 percent of their maximum potential age (80 to 100 years versus 350 to 400 years) and store only about half their potential carbon. An eighty-year-old forest today can, in most cases\u2014 barring a major disturbance such as a windstorm or insect infestation\u2014continue to accumulate carbon for at least the next two hundred years in live and dead trees and in the soil.15 Individual trees sequester more carbon the larger they grow: A forty-inch-diameter red oak (left) adds about two-tenths of an inch to its trunk diameter every year, but this new layer of biomass stores approximately the same amount of carbon as an entire sixinch- diameter tree elsewhere in the author's backyard forest. Another management option I have is reforestation: allowing an existing field to return to forest. I have begun reforestation on a small section of lawn along the edge of my property. Over the next fifteen years, this patch of regrowing forest may store as much as twenty-five times the aboveground carbon as the grassy lawn it replaced.16 Hence, reforestation has tremendous potential to sequester additional carbon on little-used pastures, agricultural fields, vacant lots, municipal fields, and small lawns on residential properties.17 There is a good reason for this potential: a site in which the trees have 32 Arnoldia 78\/3 \u2022 February 2021 been removed\u2014either recently or long ago\u2014is in a deep carbon debt because the land stores a fraction of the carbon it once stored as a forest. Energy Use Trees, of course, also have other climate-related implications for my property. Trees standing within sixty feet of my house reduce home energy expenditure and carbon emissions by cooling the house in summer and insulating it from cold winds in winter. Not surprisingly, large trees provide significantly greater energy reductions than do small trees. A thirty-inchdiameter red maple located on the west side of a house would reduce carbon dioxide emissions by almost seven-fold compared to a two-inchdiameter red maple that is similarly placed.18 One caveat is that trees, especially conifers, located on the south side of a house increase winter fuel use by blocking solar radiation; but the drawbacks are generally offset by the substantial year-round benefits of trees located on the other three sides of a house. For example, if a thirty-inch white pine was growing on the south side of my house, it would increase winter fuel use slightly, while still providing some summer cooling, resulting in an estimated 10 pounds of additional carbon dioxide emitted annually. But the same tree on the north side of the house would reduce winter fuel use\u2014and provide greater summer cooling\u2014 resulting in the reduced emissions of an estimated 335 pounds of carbon dioxide annually.19 Trees, therefore, play an important role not only in sequestering and storing carbon but also in reducing household carbon emissions. Habitat and Biodiversity Natural climate solutions can also provide important forest habitat. Trees, as they age and grow larger, provide nesting and denning sites for a host of birds and mammals.20 They create deadwood that provides food for insects and develop large crowns that supply an abundant seed source. Even scattered trees with trunks at least sixteen-to-twenty inches in diameter in an urban setting can have outsized effects on bird diversity and abundance\u2014a role that has caused researchers to describe large urban trees as \"biodiversity hotspots.\"21 Reforestation of fields and lawns can provide additional young forest habitat (when the trees are fifteen years of age or younger), an ephemeral and uncommon habitat in the northeastern United States. Several species of birds (like chestnut-sided warbler, prairie warbler, indigo bunting, and brown thrasher) and the rare New England cottontail prefer dense, low woody vegetation found in young forests, shrublands, and disturbed open woods and are generally not found in closed forests.22 Depending on how many trees are retained or regrown on a property, and where the property is located, a small parcel may serve as a green oasis in an otherwise developed environment, or as an uncommon vegetation structure in a landscape of mostly mature forest or field, or as an extension of a larger forested patch. My property best exemplifies the last scenario, as it abuts one hundred acres of contiguous forest. I frequently see and hear wood thrushes, veeries, barred owls, and pileated woodpeckers on my property. These species generally prefer mature forests or are associated with larger trees, and the wood thrush is listed as globally \"near threatened\" by the International Union of Conservation of Nature.23 Such species would almost certainly avoid my property if I converted my woods into lawn. Given that North America has lost almost 30 percent of its total bird population in the past fifty years, the natural climate solutions presented here applied across a multitude of small properties could make a real difference in stemming these population declines.24 Management for Natural Climate Solutions In general, the less I manage my property, the more climate benefits it will provide. Some tending, however, is important to allow trees to continue growing to their full potential. Lianas like the non-native oriental bittersweet (Celastrus orbiculatus), which thrive in the edge habitats characteristic of residential properties, are best cut and removed when they are growing up trees and over shrubs. Bittersweet will reduce the growth rate (and carbon uptake) and eventually kill trees by intercepting much of the sunlight in the canopy and by strangling the Backyard 33 trunk.25 The native poison ivy (Toxicodendron radicans) and grape (Vitis spp.) are generally more benign than bittersweet, but they function similarly and can proliferate in edge habitats, so I generally cut these vines at the base of my trees to give the trees every advantage to remain healthy and sequester the most carbon. With less management, tree branches inevitably grow close to my house and into my driveway and need to be trimmed periodically. After trimming, I deposit the branches in a brush pile or scatter them into the woods rather than chipping them or carting them away. Brush piles serve as cover and den habitat for a variety of small animal species such as red-backed salamanders, red-spotted newts, wood frogs, wrens, whitethroated sparrows, juncos, and box turtles.26 Trees will also die over time from insects, pathogens, and other causes and can be a hazard if houses, cars, or recreational spaces are in the fall zone. Common sense dictates that these should be cut down. But if dead trees are not a hazard, they provide considerable benefits if left standing and are not an indication that the forest is \"unhealthy\" and needs to be fixed. Though no longer sequestering additional carbon, standing dead trees continue to store existing carbon, often for decades, as the carbon is released slowly via decomposition.27 Dead trees also provide habitat for cavity-nesting birds and mammals and serve as an abundant source of insect food for woodpeckers and other barkgleaning birds like nuthatches. On my property, a standing dead elm tree (Ulmus americana) Regrowing forests can quickly store far more carbon in the vegetation than lawn grass\u2014as much as twenty-five times more in only fifteen years\u2014while also providing superior habitat. With this in mind, the author has begun a small reforestation project in an area previously maintained as lawn. Backyard 35 is used each year by a pair of yellow-bellied sapsuckers as a nest site. When I need to remove a dead tree that poses a hazard, I move it into the woods after cutting it. Similarly, when large branches and trees fall during storms, I move them off the driveway and lawn and into the woods and use some for firewood. I also resist cleaning up downed branches and trees in the woods. Downed logs serve as habitat for a host of animals, replenish nutrients and carbon to the soil, act as germination sites for new tree seedlings, and store large amounts of carbon, often for decades.28 Reforestation also requires little to no management. Tree growth is the default process in the Northeast, and the vegetation will naturally self-organize into a forest over time if a landowner simply stops mowing a lawn or field. The cessation of mowing will also add to the carbon benefits of reforestation by eliminating a significant source of emissions.29 A tall grass layer will inhibit tree growth because of competition and shading, and therefore shrubs, even thorny invasives like multiflora rose (Rosa multiflora), will generally facilitate tree seedling growth by reducing the grass layer and protecting the seedlings from deer browsing.30 In most cases, tree seedlings will eventually grow above the shrubs and reduce shade-intolerant shrub species; however, in some instances, a dense shrub layer can suppress further tree growth beneath it.31 In such cases, selectively removing some shrubs can be beneficial. Planting trees can supplement and speed up natural reforestation, but it can be expensive and labor-intensive, and is ultimately unnecessary unless a homeowner is interested in an immediate screen planting or a particular species that does not grow nearby. The Final Look Ultimately, implementing natural climate solutions is an exercise in restraint and may challenge a homeowner's sense of aesthetics. Indeed, given the choice, many homeowners prefer a relatively open, tidy property, with a few trees, long views, and unobstructed sunsets. But a property stewarded for natural climate solutions can offer a beauty not found in more open landscapes. On my property, I appreciate the delicate beams of light that pass through the foliage and columnar tree trunks in the early or later parts of a summer day; the brilliant reds, yellows, and oranges that envelop the property each autumn; and multitudes of snowor ice-covered branches on a winter day. For six months of the year, when the leaves have fallen from the deciduous trees, the views lengthen and sunsets emerge. Even during the growing season, I enjoy surprisingly long views because most of the foliage on the large deciduous trees is above rather than below the sightlines. In the small area where I have begun reforestation, sightlines are reduced and the brushy patch of tall grass, young trees, and shrubs look unkempt compared to my neighbors' adjacent, close-cropped lawn. Yet this management decision comes with other aesthetic rewards: insects busily foraging on the tall goldenrods that bloom in late summer and the flash of goldfinches and white-throated sparrows drawn to the seed source in this brushy new habitat. In the end, there is a natural beauty that accompanies the climate and biodiversity benefits of leaving more vegetation intact. Faced with runaway carbon dioxide levels and a rapidly warming climate, property owners can leverage the carbon-absorbing power of trees by keeping them standing and growing and by allowing an existing field to revert to forest by not mowing. In this way, we can play an important role in the solution by doing less and letting nature do more. Endnotes: 1 Current carbon dioxide levels are posted on https:\/\/ www.co2.earth\/. The long history of carbon dioxide levels on earth are discussed in Lindsey, R. 2020. Climate change: Atmospheric carbon dioxide. NOAA Climate.gov. Retrieved from https:\/\/www.climate. gov\/news-features\/understanding-climate\/climatechange- atmospheric-carbon-dioxide; and in Kolbert, E. 2014. The sixth extinction: An unnatural history. New York: Henry Holt and Company. 2 Harris, N. L. 2020. Young forests capture carbon quicker than previously thought. World Resources Institute. Retrieved from https:\/\/www.wri.org\/blog\/2020\/09\/ carbon-sequestration-natural-forest-regrowth 3 Adams, J. M., Faure, H. F. D. L., Faure-Denard, L., McGlade, J. M., and Woodward, F. I. 1990. Increases in terrestrial carbon storage from the Last Glacial Maximum to the present. Nature, 348(6303): 711-714. Facing page: Trees continue storing carbon long after they have fallen. Therefore, retaining logs and branches on the forest floor provides additional climate benefits and also adds new habitat types. 36 Arnoldia 78\/3 \u2022 February 2021 4 Harvard Forest. 2013. Schoolyard LTER database: Tree biomass equations. Retrieved from https:\/\/ harvardforest.fas.harvard.edu\/sites\/harvardforest.fas. harvard.edu\/files\/Tree%20Biomass%20Equations%20 2013.pdf 5 Schlesinger, W. H. 1997. Biogeochemistry: An analysis of global change. San Diego: Academic Press. 6 Stephenson, N. L., Das, A. J., Condit, R., Russo, S. E., Baker, P. J., Beckman, N. G., \u2026 and Zavala, M. A. 2014. Rate of tree carbon accumulation increases continuously with tree size. Nature, 507(7490): 90-93. 7 Data on average carbon storage in Connecticut forests was calculated using the USDA Forest Service, Forest Inventory and Analysis Program EVALIDATOR tool. Retrieved from http:\/\/apps.fs.usda.gov\/Evalidator\/ evalidator.jsp 8 Data on large tree density in Connecticut forests was calculated using the USDA Forest Service, Forest Inventory and Analysis Program EVALIDATOR tool. 9 Reinmann, A. B., Smith, I. A., Thompson, J. R., and Hutyra, L. R. 2020. Urbanization and fragmentation mediate temperate forest carbon cycle response to climate. Environmental Research Letters, 15(11): 114036. 10 Fargione, J. E., Bassett, S., Boucher, T., Bridgham, S. D., Conant, R. T., Cook-Patton, S. C., \u2026 and Gu, H. 2018. Natural climate solutions for the United States. Science Advances, 4(11): eaat1869; Foster, D., Aber, J., Cogbill, C., Hart, C., Colburn, E., D'Amato, A., \u2026 and Thompson, J. 2010. Wildlands and woodlands: A vision for the New England landscape. Cambridge: Harvard University Press; Moomaw, W. R., Masino, S. A., and Faison, E. K. 2019. Intact forests in the United States: Proforestation mitigates climate change and serves the greatest good. Frontiers in Forests and Global Change, 2: 27. 11 The emission of the entire 84.3 tons of carbon (310 tons of carbon dioxide) into the atmosphere assumes that all cut trees would be chipped and burned and none would be used for wood products. If the trees were used for wood products, approximately 20-25 percent of the tree carbon would be stored in wood. See: Nunery, J. S., and Keeton, W. S. 2010. Forest carbon storage in the northeastern United States: Net effects of harvesting frequency, post-harvest retention, and wood products. Forest Ecology and Management, 259(8): 1363-1375; United States Environmental Protection Agency. 2019. Greenhouse gases equivalencies calculator - Calculations and references. Retrieved from https:\/\/ www.epa.gov\/energy\/greenhouse-gases-equivalenciescalculator- calculations-and-references 12 Three key sources were used for this paragraph: Catanzaro, P., and D'Amato, A. 2019. Forest carbon: An essential natural solution for climate change. Amherst: University of Massachusetts; Fargione, et al. Natural climate solutions for the United States; Nunery and Keeton. Forest carbon storage in the northeastern United States. 13 Nowak, D. J. 2000, April. Tree species selection, design, and management to improve air quality. In 2000 ASLA annual meeting proceedings (pp. 23-27). Washington, DC: American Society of Landscape Architects. 14 Lu, X., Kicklighter, D. W., Melillo, J. M., Yang, P., Rosenzweig, B., V\u00f6r\u00f6smarty, C. J., \u2026 and Stewart, R. J. 2013. A contemporary carbon balance for the northeast region of the United States. Environmental science and technology, 47(23): 13230-13238. 15 Information on age and carbon storage of old growth forests in the northeastern United States was drawn from: Keeton, W. S., Whitman, A. A., McGee, G. C., and Goodale, C. L. 2011. Late-successional biomass development in northern hardwood-conifer forests of the northeastern United States. Forest Science, 57(6): 489-505; McGarvey, J. C., Thompson, J. R., Epstein, H. E., and Shugart Jr., H. H. 2015. Carbon storage in old-growth forests of the Mid-Atlantic: Toward better understanding the eastern forest carbon sink. Ecology, 96(2): 311-317. The average age of the region's forests was calculated using the USDA Forest Service, Forest Inventory and Analysis Program EVALIDATOR tool. Retrieved from http:\/\/apps.fs.usda.gov\/Evalidator\/ evalidator.jsp 16 See appendix B in: Smith, J. E., Heath, L. S., Skog, K. E., and Birdsey, R. A. 2006. Methods for calculating forest ecosystem and harvested carbon with standard estimates for forest types of the United States. General Technical Report NE-343. Newtown Square, PA: US Department of Agriculture, Forest Service, Northeastern Research Station. 17 Cook-Patton, S. C., Leavitt, S. M., Gibbs, D., Harris, N. L., Lister, K., Anderson-Teixeira, K. J., \u2026 and Griscom, H. P. 2020. Mapping carbon accumulation potential from global natural forest regrowth. Nature, 585(7826): 545-550. 18 Energy savings from trees near houses was estimated using i-Tree tools. Retrieved from https:\/\/www. itreetools.org\/tools 19 Estimated using i-Tree tools. 20 Ranius, T., Niklasson, M., and Berg, N. 2009. Development of tree hollows in pedunculate oak (Quercus robur). Forest Ecology and Management, 257(1): 303-310. 21 Stagoll, K., Lindenmayer, D. B., Knight, E., Fischer, J., and Manning, A. D. 2012. Large trees are keystone structures in urban parks. Conservation Letters, 5(2): 115-122. 23 BirdLife International. 2017. Hylocichla mustelina (amended version of 2016 assessment). The IUCN Red List of Threatened Species 2017: e.T22708670A111170926. https:\/\/dx.doi.org\/10.2305\/ IUCN.UK.2017-1.RLTS.T22708670A111170926.en. Downloaded on 13 January 2020. Backyard 37 The author standing in his backyard forest. 24 Rosenberg, K. V., Dokter, A. M., Blancher, P. J., Sauer, J. R., Smith, A. C., Smith, P. A., \u2026 and Marra, P. P. 2019. Decline of the North American avifauna. Science, 366(6461): 120-124; DeGraaf, R. M., Yamasaki, M., Leak, W. B., and Lester, A. M. 2006. Technical guide to forest wildlife habitat management in New England. Lebanon, NH: University Press of New England. 25 Webster, C. R., Jenkins, M. A., and Jose, S. (2006). Woody invaders and the challenges they pose to forest ecosystems in the eastern United States. Journal of Forestry, 104(7): 366-374. 26 DeGraaf, et al. Technical guide to forest wildlife habitat management in New England. 27 Krebs, J., Pontius, J., and Schaberg, P. G. 2017. Modeling the impacts of hemlock woolly adelgid infestation and presalvage harvesting on carbon stocks in northern hemlock forests. Canadian Journal of Forest Research, 47(6): 727-734. 28 McGarvey, et al. Carbon storage in old-growth forests of the Mid-Atlantic; DeGraaf, et al. Technical guide to forest wildlife habitat management in New England. 29 Jo, H. K., and McPherson, G. E. 1995. Carbon storage and flux in urban residential greenspace. Journal of Environmental Management, 45(2): 109-133. 30 Holl, K. D. 2002. Effect of shrubs on tree seedling establishment in an abandoned tropical pasture. Journal of Ecology, 90: 179-187; Meiners, S. J., and Martinkovic, M. J. 2002. Survival of and herbivore damage to a cohort of Quercus rubra planted across a forest\u2014old-field edge. The American Midland Naturalist, 147(2): 247-255. 31 Banasiak, S. E., and Meiners, S. J. 2009. Long term dynamics of Rosa multiflora in a successional system. Biological Invasions, 11(2): 215-224; Niering, W. A., Dreyer, G. D., Egler, F. E., and Anderson Jr, J. P. 1986. Stability of Viburnum lentago shrub community after 30 Years. Bulletin of the Torrey Botanical Club, 113(1): 23-27. Edward K. Faison is senior ecologist at Highstead in Redding, Connecticut. His work focuses on deer and moose interactions with forests, long-term forest change, and natural climate solutions."},{"has_event_date":0,"type":"arnoldia","title":"A New Look at Boston Common Trees","article_sequence":7,"start_page":38,"end_page":41,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25726","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25ea76a.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Allen, Kelsey; Oswald, W. Wyatt","article_content":"Technology changes how we see the world: think of Antonie van Leeuwenhoek's microscope or Jacques Cousteau diving with a video camera and bringing the movements of ocean life to the silver screen. For the past decade, a digital camera mounted on the roof of a ten-story building has taken photos of the Boston Common every thirty minutes. The camera is a simple consumer model, but the resulting set of photographs, numbering well over two hundred thousand, compresses time in a way that turns everyday changes within the tree canopy into meaningful patterns and trends. Within this set of images, forty seasons can be viewed as a flipbook. If you visit the Boston Common in April, you will see light-green leaves unfolding on elms (Ulmus) and the warm glow of red maples (Acer rubrum) bursting into flower, yet only in an image set like this could you determine how these hour-by-hour moments in the life of a tree correspond to seasons past. Ten years can be viewed simultaneously. Seasonal shifts can be visualized in a way that surpasses our on-the-ground experience. Moreover, thanks to image-analysis software, data can be extracted from the photographs, allowing researchers to quantify the \"greenness\" of the canopy as it changes through the growing season and from year to year. We know that global climate change is impacting plant phenology. Already, for instance, researchers have described discernable differences between flowering times for herbarium specimens that were collected one hundred years ago and those that have been collected in recent years. So far, however, the photographs of the Boston Common have shown relatively consistent leaf-out times in the spring, with the exception of 2012. The sequence of photos from that year shows the details of the springtime green-up, when anomalously warm temperatures in March triggered leaves to emerge two to four weeks earlier than other years. The elms turn green first, but not because of leaf emergence; in fact, we are seeing the maturation of samaras, the elms' winged fruits. Leaf out of the elms, along with the Common's red maples, lindens (Tilia), oaks (Quercus), and scholar trees (Styphnolobium japonicum), follows over the next few weeks. As trees on the Boston Common respond to climate change in the future, ongoing photography may reveal that years like this become less anomalous. At the other end of the growing season, the deciduous trees of the Boston Common start to prepare for winter by breaking down their photosynthetic machinery during the second half of October. The timing of those changes has not varied much over the last ten years. In the set of photos from 2018, for instance, we can see the visual transformation of the landscape that occurs each fall, with the faded greens of early autumn giving way to patches of gorgeous color, including yellow elms and reddish-brown oaks. Then, by the last week of November, the leaves have all fallen, exposing the scaffolding of branches that held them aloft all summer long. And at the tips of those branches are buds, poised to burst open in spring and start this cycle anew. Further reading Oswald, W. W. and Richardson, A. D. 2015. Tracking the seasonal rhythms of Boston Common trees. Arnoldia, 73: 36-39. Primack, D., Imbres, C., Primack, R. B., Miller-Rushing, A. J., and Del Tredici, P. 2004. Herbarium specimens demonstrate earlier flowering times in response to warming in Boston. American Journal of Botany, 91: 1260-1264. Richardson, A. D. 2019. Tracking seasonal rhythms of plants in diverse ecosystems with digital camera imagery. New Phytologist, 222: 1742-1750. Kelsey Allen is a student at Emerson College, studying literature and environmental science. W. Wyatt Oswald is a professor in the Marlboro Institute of Liberal Arts and Interdisciplinary Studies at Emerson College. He is a research associate at Harvard Forest. Boston Common Trees 41 JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER ALL PHOTOS BY W. WYATT OSWALD Each column shows the Boston Common during the first week of the month\u2014revealing differences year over year."},{"has_event_date":0,"type":"arnoldia","title":"Case of the Anthropocene","article_sequence":8,"start_page":42,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25727","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25eab6d.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"On December 18, 1994, three cave explorers squeezed into an opening of a cliff overlooking the Ard\u00e8che River in southern France. At the back, a whisper of cool air prompted them to prize stones from a narrow passage and worm forward headfirst. After ten feet, they encountered a thirty-foot drop into a large chamber. Beneath them, as it turned out, the cave walls were covered with paintings. Some appeared almost fresh. First, the explorers found a mammoth drawn in red pigment, then woolly rhinoceroses, cave lions, and compositions made entirely of human handprints. Researchers would later determine that a landslide sealed the main entrance to the cave, now known as Chauvet Cave, twenty-eight thousand years ago, safeguarding hundreds of paintings and wall engravings. Eighteen thousand years later, glaciers had retreated from much of Europe, and many of the animals depicted in Chauvet Cave had gone extinct. Humans in Mesopotamia were domesticating wheat and barley. Fast forward another nine thousand years to the completion of the first recorded circumnavigation of the globe in 1522. Eventually, in the summer of 1833, an English sailing ship departed London, bound for Australia. On the upper deck, the captain diligently monitored two sealed glass cases planted with ferns, grasses, and mosses. About six months later, the ship arrived in Sydney with all but three of the plants still alive. The case was opened only once; moisture cycled naturally inside the enclosure. On the return trip, the cases were packed with ferns that survived air temperatures fluctuating between twenty and more than ninety degrees Fahrenheit. In fact, the cases were so effective that stowaway seeds germinated in the soil. A shipment of plants between the antipodes might seem like a minor historical footnote, but in a new book, The Wardian Case: How a Simple Box Moved Plants and Changed the World, historian Luke Keogh describes the shipment as a profound inflection point in the history of the Earth. Keogh first became interested in these enclosed glass cases while curating an exhibit at the Deutsches Museum in Munich. The exhibit opened in 2014 and focused on the Anthropocene, a term for our current geologic era that acknowledges the enormity of humancaused environmental change. Millions of years from now, our present moment will appear in the geologic record as an abrupt transition char- Case of the Anthropocene Jonathan Damery Luke Keogh. The Wardian Case: How a Simple Box Moved Plants and Changed the World. The University of Chicago Press and the Royal Botanic Gardens, Kew, 2020. BIODIVERSITY HERITAGE LIBRARY\/FROM US BUREAU OF ENTOMOLOGY (1913) BULLETIN NO. 120 DAMERY, J. 2021. CASE OF THE ANTHROPOCENE. ARNOLDIA, 78(3): 42-43 Wardian cases moved plants around the world, along with insects and other organisms. Book Review 43 acterized by rapid climate change, sea-level rise, and mass extinction\u2014an imprint far more permanent than the markings at Chauvet Cave. The unprecedented biotic exchange ushered in by the experimental plant shipments between London and Sydney is a piece of this story. The experiments had been orchestrated by an affable English physician named Nathanial Ward and the nurseryman George Loddiges. Previously, it had been exceptionally difficult to ship live plants over such long distances. In addition to the general perils of sea travel (salt spray, tempestuous weather, foraging rodents), fresh water was a scarce resource and could seldom be spared for plants. In a backyard experiment, Ward discovered that plants could be sustained within an enclosed glass container for long periods without supplemental water. When such cases were used aboard ships, they solved many of the persistent problems associated with long-distance plant transport. In a follow- up experiment in 1834, Ward sent six cases to Egypt and Syria, and when the plants were received, scarcely a leaf was reported missing. Keogh follows the Wardian case as it became a commonplace tool, not only for moving botanical curiosities but also for transporting crops (including tea, Camellia sinensis, and rubber, Hevea brasiliensis) that supported the endeavors of Western empire-building. Also, because Wardian cases contained soil, the plants invariably arrived with insects and pathogens in tow. \"To move plants was to move ecosystems,\" Keogh writes. Some of these newcomers proved devasting, including coffee rust (Hemileia vastatrix), which erupted in Ceylon (now Sri Lanka) in 1869 and subsequently decimated plantations in many coffee-growing regions around the world. Altogether, this global churning\u2014which continues in a post-Wardian world\u2014accumulates to dramatic effect. Keogh, for instance, cites a study suggesting that approximately nine out of ten invertebrate pests in the United Kingdom arrived on live plants. Certainly, the Wardian case was just one innovation within the broader scope of the Anthropocene. The case gained traction at a moment of enormous industrialization and fossil fuel use. While the first Wardian cases were transported on sailing ships, steam power soon predominated. Carbon dioxide levels in the atmosphere would mount. Moreover, industrial agriculture favors monocultures, which are especially susceptible to pests and pathogens (like coffee rust) that spread rapidly in the Wardian era. In a curious twist, Keogh recounts how, in the early twentieth century, entomologists used Wardian cases to intentionally transport insects to control invasive plants and other pests that had been imported in earlier shipments. By the 1920s, plant quarantines and import restrictions slowed the use of Wardian cases, but it was the airplane that finally rendered them obsolete. Now live plants can be moved without soil, wrapped in plastic, and mailed directly to inspection sites before being admitted into a country, assuming importers follow the rules. Yet pests and pathogens continue to spread. The emerald ash borer (Agrilus planipennis) was first identified in the United States in 2002 and likely arrived burrowed within wood shipping materials. The Asian longhorned beetle (Anoplophora glabripennis) arrived in a similar fashion before 1996. In this light, the Wardian case was only one contributor to this dramatic biotic exchange. Not only has the admixture continued to the present but humans began moving plants long before Nathaniel Ward arrived on the scene. Ward's main innovation, Keogh stresses, was the enclosed system. Also, not insignificantly, Ward was a charismatic individual who used his social connections to promote the case. For Ward, awaiting news on his inaugural shipment to Australia, the long-term implications of his cases would have been impossible to imagine. Thinking about consequences two hundred years in the future is almost beyond the realm of comprehension\u2014almost as unlikely as the painters at Chauvet Cave imagining researchers studying their work more than thirty thousand years later. Yet the concept of the Anthropocene asks us to think even further ahead. In 1833, the captain of the ship to Australia penned a congratulatory letter to Ward: \"Your experiment for the preservation of plants alive \u2026 has fully succeeded.\" The case of the Anthropocene challenges us to reconsider the meaning of our own small successes. Jonathan Damery is the editor of Arnoldia."},{"has_event_date":0,"type":"arnoldia","title":"Planting Edo: Pinus thunbergii","article_sequence":9,"start_page":44,"end_page":45,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25728","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d25eaf25.jpg","volume":78,"issue_number":3,"year":2021,"series":null,"season":null,"authors":"Saunders, Rachel","article_content":"In February 2020, we opened our largest ever exhibition at the Harvard Art Museums, never anticipating that, a month later, the doors of the museums would close due to the pandemic. Painting Edo: Japanese Art from the Feinberg Collection features 120 paintings arranged as an immersive, in-person experience. At the onset of the closure, when I rushed about my office gathering books and papers, I expected to be away for only a few weeks, but as our exile from the galleries continued, we adapted to virtual close-looking through an online exhibition and Zoom events. What I hadn't realized was how significantly this new form of looking would alter my own vision of Edo painting. One work that I came to see differently was Old Pine by the eighteenth-century painter Ito\u02c9 Jakuchu\u02c9 . It is by no means a fresh observation that artists of the Edo period (1618-1868) were extremely interested in the natural world. Jakuchu\u02c9 is celebrated today for the magical hyper-realism of his polychrome paintings of flowering plants, aquatic animals, and especially chickens, which he is said to have kept so that he could observe the complexity of their feathers daily. Old Pine, by contrast, is executed in gestural monochrome ink. The painting is modestly sized, but the radical proximity from which the tree is painted\u2014so close that it cannot be contained within the picture plane\u2014makes an encounter with it feel as overwhelming as standing beneath an enormous conifer. Pines have a long history in East Asian art and are among the primary subjects of ink painting. In the vocabulary of this spare, highly intellectualized mode of painting, pines represent resilience, longevity, and the integrity of the upright scholar-gentleman. Identification of a painted tree as \"a pine\" is all that is sufficient to trigger these associations, since ink painting valorizes capturing the essence of a thing over mere verisimilitude. Jakuchu\u02c9 had clearly captured an individual arboreal essence, but it was not until a botanist's eye was turned upon it that the true level of Jakuchu\u02c9 's observation emerged. With Zoom, the distance between the painted plants in the galleries and their living counterparts at the Arnold Arboretum melted away. This enabled a new privilege of simultaneously looking at living and painted plants with the Arboretum's Michael Dosmann and Ned Friedman. Our conversations led to a series of public virtual events. With this botanical view, the eccentrically angled branches, plated bark, and textured twigs of Jakuchu\u02c9 's \"pine\" resolve almost immediately into features of a \"black pine,\" or Pinus thunbergii (kuromatsu in Japan). When we view the painting, a major limb\u2014 covered, dragon-like, in scaled bark\u2014thrusts up from the bottom left-hand corner, only to disappear beyond the right-hand border. It curves back into the frame at the top right, from where an angular branch, brushed in several switchback strokes, descends. This dramatically contorted form echoes the Japanese black pines growing at the Arnold Arboretum (see accession 11371), and so, too, does the orientation of the painted needles: spiky lateral marks from a wide brush that flare from axial twigs. But the precision of Jakuchu\u02c9 's observation is evident beyond these most prominent elements. A variety of lichen-like dots peppers the branches, the largest pressed from the side of an inked brush, and the smaller nubby marks from its tip. What I had read as an anomalous abundance of mosslike texture strokes, Ned's eye revealed as the closely observed characteristic texture of black pine twigs, formed by the unusual persistence of bracts, which can remain for up to two years after their sets of paired needles fall. In an inscription brushed in 1755, Jakuchu\u02c9 wrote: \"Flowers, birds, grasses, and insects each have their own innate spirit. Only after one has actually determined the true nature of this spirit through observation should painting begin.\" Old Pine shows just how thoroughly Jakuchu\u02c9 took this dictate, not only in his obsessively observed and painstakingly detailed polychrome paintings but also, we can now see, in the spare and immediate genre of ink painting. Rachel Saunders is the Abby Aldrich Rockefeller Curator of Asian Art at Harvard Art Museums. Planting Edo: Pinus thunbergii Rachel Saunders"},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25697","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270a76a.jpg","title":"2021-78-3","volume":78,"issue_number":3,"year":2021,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Pandemic Digitization","article_sequence":1,"start_page":2,"end_page":4,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25714","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24eaf6e.jpg","volume":78,"issue_number":2,"year":2020,"series":null,"season":null,"authors":"Brach, Anthony R.","article_content":"Listening to Vivaldi's Four Seasons, I began my day transcribing data from herbariumspecimen labels. The melodies and the early morning light mixed, and I entered the zone, my fingers typing rhythmically with the music. Staff at the Harvard University Herbaria transitioned to working from home due to the coronavirus pandemic on March 16. In the early weeks of this new routine, I was transcribing detailed data from specimens collected in Wyoming\u2014locations like Devils Tower and Yellowstone\u2014but instead of handling the physical specimens, I was working from images on my screen. This current work has been very different from the normal day-to-day curatorial activities at the Herbaria. Researchers, who we would normally be assisting, have been unable to visit the collections. Our team, likewise, was initially unable to be on-site for routine activities like processing incoming and outgoing shipments of loans, gifts, and exchanges. We could not mount new specimens or file them into the collection; nor could we update specimens with new taxonomic determinations. We have even discouraged other institutions from sending materials given potential shipping and handling delays. Before this began, however, our team was busy with a long-term effort to share images and data from our collections online, and this meant we could use the same images to continue digitization projects remotely as well. Over the past 170 years, the Herbaria have amassed more than five million specimens, making our collections one of the largest in the world. Given the scale, specimens have been digitized on a project-by-project basis. About one-quarter of our total holdings have been digitized to date. I like to think of imaging and transcription of specimen labels as \"publishing\" unfinished symphonies composed by botanists. Without digitization, their collections are often hidden in the Herbaria, requiring either in-person visits or potentially risky shipments of specimen loans. Among the Wyoming specimens, for instance, I enjoyed databasing those collected by Reed Rollins, a Harvard professor and longtime director of the Gray Herbarium. Many of his extensive collections of the mustard family (Brassicaceae) were redetermined by his student Ihsan Al-Shehbaz, who followed Rollins as the world's foremost taxonomist of this family. Now digitized, their collaborative work has become available for study by a new generation of researchers. Before the pandemic, our curatorial team was in the middle of three collaborative digitization projects funded by the National Science Foundation and coordinated through the Thematic Collection Network. One focuses on the Southern Rockies. The second focuses on the vascular flora of the South Central United States, particularly Texas and Oklahoma. The third is called Endless Forms (or Plants on Edge) and focuses on fifteen families of rare and endangered plants with unique morphological adaptations, including orchids (Orchidaceae), cacti (Cactaceae), and sedums (Crassulaceae). Combined, these projects include about 470,000 specimens. Our director of collections Michaela Schmull and the director of informatics Jonathan Kennedy have orchestrated our curatorial team's digitization efforts so that, rather than pulling collections piecemeal by individual states (states are filed alphabetically for each species), all vascular plants from the United States and Canada were added to the queue. This expansion (another 1.6 million specimens) is part of the Herbaria's effort to digitize the entire collection. When the closures began, I had been photographing Lupinus in the legume family (Fabaceae), and recently, I had photographed specimens of a few other families with great diversity in the Rockies, including the mustards (Brassicaceae) and saxifrages (Saxifragaceae). Now, working from our homes as a team, the thirteen of us curatorial assistants had the opportunity to loop back and record detailed Pandemic Digitization Anthony R. Brach \u222b data from specimens we had already photographed. This data entry allows the specimens to be searchable using details like the collector's name and collection date. Our team completed transcription from available images from project-related states (about 66,670 specimens) after the first couple of months of the pandemic. This could have taken three times longer if not for our work-from-home efforts. Next, we moved onto other states and provinces not part of the projects. For this second phase, I selected New York. I was born in Rochester, and when I was just a kid, I carried Peterson's Field Guide to Wildflowers of Northeastern and North-Central North America on hikes with my dad, who was an avid, knowledgeable amateur botanist. He took my brothers and me to regional parks and to the Adirondack Mountains. In my college years, I explored the Hudson River Valley and Long Island Sound, and my graduate research on the ecology of forest-understory herbs and ferns brought me back to the Adirondacks. Transcribing specimen labels for this familiar flora allowed me, in some sense, to revisit these ecosystems. Because many labels were from the nineteenth to early twentieth centuries, only a fraction were typed, while many were handwritten and of various degrees of legibility. Since the beginning of the pandemic, our team has been communicating via Slack, a chatting tool that we have used for asking questions and helping one another decipher illegible handwriting on labels. I was amused by two handwritten labels for collections from Irondequoit Bay (Rochester area) and Taughannock Falls (north of Ithaca). If not for my familiarity with these places, I do not know if I could have deciphered them. We also have a very large collection of specimens from New York that were collected by Asa Gray, the first director of the Gray Herbarium, whose handwriting has always been challenging to read. I was fascinated to see specimens collected more than one hundred years ago near my hometown and from other familiar places. In 1889, collector John Dunbar told Charles During the pandemic, curatorial staff at the Harvard University Herbaria have been recording collection data from previously photographed herbarium specimens. Often the intrigue is in the details. THE HERBARIUM OF THE ARNOLD ARBORETUM AND THE GRAY HERBARIUM, HARVARD UNIVERSITY 4 Arnoldia 78\/2 \u2022 November 2020 Sprague Sargent, the director of the Arnold Arboretum, about many hawthorns (Crataegus) near Rochester that did not match any described species. Dunbar and others\u2014including his coworkers Calvin C. Laney, Henry T. Brown, and Berhard H. Slavin (all from the Rochester Parks Department)\u2014collected hundreds of specimens for Sargent. Beyond the familiar locations, some specimens included details that made these places come alive with activity. I came across a 1905 label, for instance, which noted that small boys filled their pockets with fruits from a scarlet hawthorn (Crataegus pedicellata). Others documented landscapes that were changing like the tempo of Grieg's \"In the Hall of the Mountain King\" (made popular in Fantasia). A dramatic 1907 label for another hawthorn (C. brainerdii), for instance, marked history: \"Prof. Sargent you will notice that I have changed this No. as I told you my No. 2415 was blown up by the Barge Canal work,\" a physician-botanist named Joseph V. Haberer wrote. Strikingly, the label recorded an instance of the widening of the Erie Canal, between 1905 and 1918, for use by large barges. When I encounter multiple specimens from the same collector, I often look up the person's backstory. Collections by botanists who happened to be medical doctors often catch my attention, especially since one of my sons serves as a doctor of osteopathic medicine and his brothers study pharmacy and medicine. In addition to Haberer (and Asa Gray, who trained as a physician), other medical doctors who collected specimens in New York included Henry P. Sartwell, George Thurber, Peter D. Knieskern, George G. Kennedy, and Edwin H. Eames. Their collections, too, have now been digitized for continued studies. Starting on June 15, our team transitioned to a hybrid work model, which allowed for limited entries into the Herbaria for a set number of hours, one day per week. This required strict adherence to the university's guidelines, safety protocols, and weekly coronavirus testing. It was a relief to be back but strange returning to a near-empty place, devoid of researchers. With this on-site day each week, I aimed to take \u222b care of essential services for the collections, in coordination with others during their allotted times at the Herbaria. I processed incoming shipments (after freezing to prevent potential insect problems), checked insect traps (each of the curatorial staff has an area to monitor), and photographed specimens as requested by botanists for their remote studies. I attached barcodes to a new set of two hundred herbarium sheets of Lupinus and photographed them for digitization from home. I finally reached Lupinus texensis, the brightly colored, bluebonnet of Texas. The university has encouraged staff to continue working from home, so transcription will continue to keep everyone busy. During our remote work so far, from mid-March until mid- September, our team has digitized 135,333 specimens, bringing the total number of digitized North American specimens in the Herbaria to nearly one million. These data and images can be found using the search interface on the Harvard University Herbaria website. Our team also learned how to use the Geo-Locate Project's collaborative georeferencing tool to add mapping coordinates whenever possible, starting with localities in the Southern Rockies. Throughout the pandemic, as I've been working with these digital specimens, my wife, Ying, has also been working from home. She is a forest ecologist by training. In the early months, when we left the house for walks in our neighborhood and local conservation areas, we were encouraged by the sights and sounds of spring. Plants flowered and produced leaves as usual, and the seasons have continued to flow like Vivaldi's melodies. This ceaselessness is echoed in our preserved herbarium specimens, each of which documents a particular moment from seasons past. Seasons and generations accrue. When brought together\u2014and shared with researchers and teachers\u2014the long-hidden symphonies, at last, resound. Anthony Brach is a senior curatorial assistant at the Harvard University Herbaria and a research associate of the Arnold Arboretum. Previously, between 1993 and 2012, he served as an editor of the Flora of China, while based at the Harvard University Herbaria as a Missouri Botanical Garden staff member, after completing his PhD in environmental and forest biology at the SUNY College of Environmental Science and Forestry."},{"has_event_date":0,"type":"arnoldia","title":"Redefining \"Remote Fieldwork\"","article_sequence":2,"start_page":5,"end_page":7,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25715","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24eb326.jpg","volume":78,"issue_number":2,"year":2020,"series":null,"season":null,"authors":"Hruska, Amy; Komatsu, Kimberly","article_content":"During the summer, the Smithsonian Environmental Research Center (SERC), in Edgewater, Maryland, is typically buzzing with activity. Scientific staff and volunteers arrive early in the morning to load gear into field vehicles and begin long days of research on land or at sea. Much of this fieldwork, near or in the Chesapeake Bay, deals with pressing and complex environmental threats, such as climate change and invasive species. Roughly two dozen visiting undergraduate researchers move into campus dormitories during the summer and join the research labs where they diligently work to complete independent projects in just ten weeks. In SERC's Ecosystem Conservation lab, we investigate how ecosystems respond to global threats, such as nutrient runoff, land-use conversion, and invasive species. Our plan for the summer of 2020 was to revisit over a dozen forest fragments in the Chesapeake Bay watershed for the first time in more than forty years to assess how land-use change has affected plant and songbird populations. Undergraduate researchers were to be instrumental in resurvey efforts and would have the opportunity to design complementary field experiments or surveys that would broaden their experience. Over the winter, we assembled an all-star team: Skye Austin, a rising sophomore from Shenandoah University, enthusiastic about the environment and conservation and ready for her first research experience. Rachael Brenneman, a rising senior at Eastern Mennonite University, eager for the chance to design and implement her own field research after conducting class research projects. And Julia Smith, a recent graduate of the University of Chicago and a data modeler, excited to get outside and experience the nuances of ecological field research. During any given field season, we anticipate that not everything will go as planned\u2014an unexpected storm may shift the schedule or cause extensive damage to a site, or we might add measurements to account for new field observations. This year, however, the very idea of conducting fieldwork and mentoring students seemed to hang in the balance as the coronavirus pandemic led to nationwide shutdowns and internal policy changes. As stay-at-home orders went into place in March, it was unclear how field research programs would proceed\u2014if at Redefining \"Remote Fieldwork\" Amy Hruska and Kimberly Komatsu 6 Arnoldia 78\/2 \u2022 November 2020 all. Overnight, SERC's research campus became an unrecognizable ghost town as most of the staff began to telework and only pre-approved, essential staff (including members of our lab) came in to maintain critical operations and experiments. Over time, it became clear that this would be the new normal, and as a result, the organizers of the undergraduate research program decided to take everything remote. As our lab began planning a remote field season that did not involve a plane ride, we initially inventoried existing datasets related to plant mutualisms, biodiversity, and ecosystem function, and generated a list of possible questions that undergraduate students could address while living at home, turning a fun field-based research experience into ten weeks in front of a computer screen gathering data from the web or navigating the world of statistical analyses. While this type of experience would certainly be valuable for many students, the idea of a computer-based internship did not meet the goals of our three undergraduate researchers who were eager for the chance to design and conduct field experiments. Cue inspiration from none other than Charles Darwin. While most of us go through school associating Darwin with his voyage on the HMS Beagle and the theory of natural selection, many of his theorytesting experiments took place from the comforts of his own home (see Darwin's Backyard: How Small Experiments Led to a Big Theory by James Costa). We asked, would it be possible for our undergraduate students to conduct field experiments at their family homes? Before the undergraduate researchers started in mid-June, we determined their locations in relation to SERC, their indoor and outdoor spatial constraints for an experiment, and compiled topics and resources that would help shape the type of questions they'd be able to ask. Coincidentally, everyone lived within three hours of SERC, so with extra steps to keep materials disinfected and acquire administrative approval, we could drive materials to their homes. Furthermore, everyone had outdoor space in their family yards to set up an experiment. Thus, a summer of backyard ecosystemconservation research began. Our undergraduate researchers hit the ground running. With minimal direction other than the compiled topics and resources related to our broad research themes and the agreed-upon spatial constraints, they worked together to develop an overarching research question and experimental design that they could each have in their yards. Over two weeks, they read the scientific literature and met daily to settle on one overarching question: how does nitrogen pollution from runoff affect plant and soil communities? To address this question, they would each set up sixty one-gallon pots in their yards, each pot containing two native plants. Plants within a pot could be one of three native species: Joe-Pye weed (Eutrochium purpureum), sensitive partridge pea (Chamaecrista nictitans), or Virginia wild-rye (Elymus virginicus). All possible combinations were represented, meaning that a pot could be planted with either two of the same species or two different species. Next, the team identified measurements that would allow them to answer more specific questions based on their individual interests. Skye was interested in the capacity of these native plants to uptake added nitrogen under different diversity treatments. Rachael asked how added nitrogen and plant diversity treatments affect the soil microbial community. And Julia wanted to understand how nitrogen addition and diversity treatments affect plant competition. Everyone was responsible for collecting the data that would be needed to address each of these three questions. They would take plant growth measurements, collect soil and invertebrate samples, and harvest plants for analyses of biomass and nitrogen content. After settling on the questions, experimental design, measurements, and materials, we spent a week purchasing and preparing all the required materials. We then made a ten-hour road trip to drop off the materials at each house. Traditionally, lab mates would help with project setup, but this year, the undergraduate researchers were left to handle those steps on their own. Previous page: Julia Smith, an undergraduate researcher at the Smithsonian Environmental Research Center, receives a delivery of remote research supplies. PHOTO BY AMY HRUSKA Remote Fieldwork 7 To ensure each researcher made the same judgment calls during setup (such as how to orient the plants in the pot), they held a multi-hour video meeting to discuss the process. Later, long video discussions became a reoccurring theme as the team took each measurement for the first time and harvested their plants at the end of the experiment. But various household members (parents and friends) did help each student with the setup (and maintenance and harvest). In some cases, parents became just as invested in the success of the plants as the undergraduate researchers themselves, checking on the experiment periodically just to see how the plants were getting along. In total, the experiment ran for a little over five weeks, with plants exposed to four weeks of fertilizer treatments in concentrations that matched those found in runoff from residential yards. After the last plant was harvested, we made a final road trip to collect their samples, as well as the equipment loaned for the summer. Back at SERC, we dried and stored samples that will be analyzed in the lab at a later date. For our undergraduate researchers, a final virtual presentation bookended their summer experience. Together, the researchers eloquently presented their fieldwork experience and discussed how they designed a single experiment to answer a host of meaningful questions related to ecosystem conservation. While this summer was a far departure from our initial plans, and a deviation from what is traditionally considered remote fieldwork, each undergraduate researcher experienced the hallmarks of conducting field research. Everyone coped with the heat and humidity of the DC, Maryland, and Virginia metropolitan area as they took their late-summer measurements. They anxiously sat and watched their pots from indoors as Hurricane Isaias brought heavy winds and rains to their yards. They all agreed to add herbivory observations to their data collection after each experiment had evidence of unintended interactions with residential wildlife. But, most importantly, everyone felt the ownership and satisfaction that can only come from developing and completing an experiment. Data analysis for this project is ongoing and will continue through the fall and winter. Many of the samples still need to be processed in the lab to determine plant biomass, and leaf and soil nitrogen content. While Julia is currently starting her doctorate, Skye and Rachael have continued as fall interns in the Ecosystem Conservation lab, working to finish these analyses and lead the efforts to publish their results. The initial results are beginning to tell an exciting story as to how plant diversity may help combat nutrient pollution. The data suggest that some species can continue to grow just as well under high nitrogen conditions from runoff and in different diversity treatments. A bonus of conducting remote research from home this summer was the realization that undergraduate researchers can, in some cases, continue to be supported once they return to school. As our lab continues to function over video conferencing, undergraduate researchers can be involved in lab meetings and SERC virtual events. And as SERC moves through the phases of its reopening plan as coronavirus cases drop in the region, the undergraduate researchers will finally be able to make it into the lab to process their samples. The current pandemic has changed many aspects of our day-to-day lives and how we conduct science. At times, these changes are overwhelming and do not have clear resolutions. Yet, this pandemic has also demonstrated our ability to be resilient and adapt to the previously unimaginable. Our ability to pivot from an in-person field program to conducting remote science in our backyards is one of many examples of how field scientists have coped this summer. These examples, however, should not come as a surprise. As field scientists, we know that disruptions are inevitable, and we need to be flexible and open to new solutions. If anything, conducting science during an unprecedented time is what field research has been preparing us for all along. Amy Hruska is a postdoctoral fellow in the Ecosystem Conservation lab at the Smithsonian Environmental Research Center (SERC). Kimberly Komatsu is the senior scientist and principal investigator of the Ecosystem Conservation lab at SERC."},{"has_event_date":0,"type":"arnoldia","title":"Closing the Book on Sargent's Weeping Hemlock","article_sequence":3,"start_page":8,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25716","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24eb36a.jpg","volume":78,"issue_number":2,"year":2020,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Sargent's weeping hemlock (Tsuga canadensis 'Sargentii') is one of the world's greatest dwarf conifer cultivars in terms of its beauty, longevity, and stability. As opposed to the typical eastern hemlock with a tall straight trunk reaching upwards of a hundred feet, the weeping variety is a totally horizontal tree that can form a giant dome of foliage up to twenty feet high by forty feet across\u2014\"a vernal fountain of perpetual joy\" is what one writer called it.1 The tree was discovered in the mid-nineteenth century in the Hudson Highlands,2 about sixty miles northeast of New York City. This part of the world was a critical supply depot for the Continental Army during the American Revolution, and later its scenic vistas inspired both the Hudson River School of painting and the \"picturesque\" landscape movement championed by Andrew Jackson Downing. This region is one of the areas where modern American ornamental horticulture first took root, and many of its earliest practitioners built country estates in the area based on aesthetic principles that Downing laid out in his writings from the 1840s and 50s. One such horticultural pioneer was Henry Winthrop Sargent, the man for whom the weeping hemlock was named. In 1841, he purchased a twenty-two-acre parcel of woodland overlooking the Hudson River at Fishkill Landing\u2014also known as Fishkill-on-the-Hudson\u2014where he developed a country estate called Wodenethe, which included sweeping vistas and an especially notable collection of conifers.3 Sargent's younger cousin Charles Sprague Sargent, the first director of the Arnold Arboretum, would describe the conifer collection as \"the most complete in the United States.\" H. W. Sargent made his place famous by describing the design and construction of its grounds in the supplement to the sixth edition of Downing's classic book, A Treatise on the Theory and Practice of Landscape Gardening, published in 1859. In an update to the supplement, in 1875, Sargent produced a vivid description of the \"gardenesque\" landscape effects he sought to achieve through the use of exotic plants with extreme growth habits and foliage textures and tints. \"There should be certain groups all color, other groups all form, and others again pendulous or drooping,\" Sargent wrote. \"But these colors and forms must be harmoniously arranged by very careful blending. Sometimes in contrast (not so great as to shock), and sometimes by the delicate merging and intermingling of one color with another, the deeper and darker first, to disappear and melt away as it were into the lighter and fairy-like tones.\" For Sargent, landscape gardening was more about art than science, and the garden itself was a kind of living sculpture. The weeping hemlock that now bears his name fit so perfectly into Sargent's gardenesque landscape style that, as has been said, had he not introduced it, he would have invented it.4 My own interest in Sargent's weeping hemlock began in 1970. I had just moved to Boston from California and was teaching biology to children ages five through eighteen at an experimental school in Watertown, Massachusetts. Always on the lookout for interesting field trips, I visited the Arnold Arboretum for the first time in the fall of that year. In my aimless wandering, I came across a bizarre, low-growing tree with twisted, ribbon-shaped branches, the likes of which I had never seen before\u2014it was Sargent's weeping hemlock. Why did it have its amazing shape? Where did it come from? How did it get here? Although I did not recognize it at the time, I had been seduced by the tree and the Arboretum where it was growing. Closing the Book on Sargent's Weeping Hemlock Peter Del Tredici Facing page: The story of Sargent's weeping hemlock often centers on the plant's namesake, Henry Winthrop Sargent, who grew the horticultural curio at his estate, Wodenethe, in the Hudson River Valley. But archival discoveries have introduced new characters to the story. SARGENT\/ARNOLD ARBORETUM ARCHIVES; WODENETHE (DOWNING, 1859)\/BIODIVERSITY HERITAGE LIBRARY; ARNOLD ARBORETUM SPECIMEN (15820*B )\/JONATHAN DAMERY; DUTCHESS COUNTY, NY, MAP (1850)\/ LIBRARY OF CONGRESS, GEOGRAPHY AND MAP DIVISION \u222b 10 Arnoldia 78\/2 \u2022 November 2020 When I finally began working at the Arnold some nine years later, in 1979, my interest in Sargent's weeping hemlock was rekindled when Augustus M. Kelley, publisher of Theophrastus Books in Little Compton, Rhode Island, wandered into the Dana Greenhouses where I was the assistant plant propagator and, without introducing himself, started talking to me about weeping hemlocks. At some point in the conversation, after I had expressed interest in the topic, I mentioned that I had a theory about why hemlocks weep. Gus said he'd like to hear it and, after listening carefully for about five minutes, asked if I would write it up. I told him I'd think about it, and a year later, I published my first-ever article for Arnoldia, the magazine of the Arnold Arboretum: \"Sargent's Weeping Hemlock Reconsidered.\" As soon as the article was published, however, I discovered several new references related to the origin of the tree, including one that pushed its first mention in print from 1875 to 1868. Gus suggested that this new information warranted an update of the weeping hemlock story and offered to publish a book about the tree if I would write it. To make a long story short, A Giant Among the Dwarfs came out in 1983, providing a new account of the tree's history. There were still gaps in the story, of course, but I did my best to fill them with well-reasoned speculation. Predictably, after the book came out, people wrote to me with new information about various specimens of Sargent's weeping hemlock, which I dutifully stashed away in a file folder, never really expecting to revisit the subject. In the years since the publication of my book, the internet was invented, and the door that I had closed some thirty-six years ago cracked open with the unexpected discovery of a statement from H. W. Sargent himself, in 1880, about who actually discovered the tree that carried his name. One thing led to another, and the cold case of the true discoverer of Sargent's weeping hemlock suddenly got very hot. With the help of various websites\u2014especially the Biodiversity Heritage Library\u2014I was able to access a slew of old references that shed new light on the story of how this sublime conifer came into being. And so, it is with some trepidation that I make my third attempt at resolving the contradictions that have plagued Sargent's weeping hemlock since its discovery. Hopefully, this time will be the charm. In the Beginning Based on research that I completed for A Giant Among the Dwarfs, I concluded\u2014correctly as it has turned out\u2014that the first written reference to Sargent's weeping hemlock was from 1868. The critical passage appears in The Book of Evergreens by Josiah Hoopes, a well-known nurseryman and conifer specialist, in the midst of his description of H. W. Sargent's Wodenethe estate. \"Near the mansion are two very handsome specimens of Araucaria imbricata, grown in boxes,\" Hoopes wrote, referring to the monkey puzzle tree (now A. araucana). \"These had attained the height of 5 or 6 feet, and were perfect examples of this species in a young state. Near these we noticed a remarkable variety of the Hemlock Spruce, of dwarfish habit, with long drooping branchlets, and altogether quite unique in character. This plant was found growing on the mountains near by.\" On the basis of the description alone, one could not say absolutely that Hoopes was talking about Sargent's weeping hemlock, but when the location of the discovery on a nearby mountain is added, the plant could be nothing else. Hoopes, nonetheless, omits the tree from the main body of the book where the \"hemlock spruce\" (listed as Abies Canadensis5) and two of its varieties are discussed, suggesting that the plant was relatively unknown in 1868. I recently found a second reference to the plant at Wodenethe in an 1874 article about mutant conifers by one Thomas C. Maxwell, a nursery owner from Geneva, New York. \"On Mt. Hounes, Fishkill-on-the-Hudson, is found a sport from our well known Hemlock,\" Maxwell reports. \"The species we all know is remarkably graceful and beautiful, lofty and grand, but this sport grows down as persistently as the Kilmarnock Willow\u2014a real deformity, and yet on Mr. Sargent's lawn it is one of the most interesting and ornamental plants in his entire collection\u2014 'a thing of beauty,' with which scarcely another tree or plant on these most beautiful grounds or in all the land can compare.\" It took me a while to figure out that \"Mt. Hounes\" was an alternate spelling for what is today known as Honness Mountain, a 906-foot \"peak\" near the present-day town of Fishkill\u2014 about five miles northeast of Wodenethe.6 Maxwell's description of the tree is particularly noteworthy because he describes how the wild weeping hemlock that was discovered on Honness Mountain\u2014\"a real deformity\"\u2014was transformed into \"a thing of beauty\" after being cultivated at Wodenethe, as if the plant had somehow gone to finishing school. Sandwiched between these two early references to H. W. Sargent's stunning new hemlock was a more complete description of the tree published by Frank Jessup Scott in his monumental work, The Art of Beautifying Suburban Home Grounds of Small Extent. Curiously, there are two different versions of this book with an 1870 publication date: One is 274 pages long and deals mainly with garden design issues. The other contains an additional 244- page section titled \"Part II: Trees, Shrubs and Vines,\" which contains detailed descriptions of woody ornamental plants suitable for planting in home landscapes. In the shorter of the two 1870 editions, Sargent's weeping hemlock is mentioned only in the fifteenth chapter, \"Plans of Residences and Grounds.\" This section of the book presents written descriptions of twenty-nine hypothetical landscape layouts, along with detailed drawings showing the locations of recommended plants. In the seventh plan (as well as in seven others7), Scott uses the letter H to designate the position of a plant he identifies as \"Sargent's hemlock, Abies canadensis inverta\" and recommends that \"its main stem to be kept tied to a stake until it has a firm growth six feet high.\" Remarkably, this first attempt at giving Sargent's weeping hemlock a proper scientific name is one of only two times that the epithet inverta appeared in print. In the longer of the two 1870 editions of Suburban Home Grounds, which is identical in all respects to an 1873 edition (except for the date), Scott preserves the use of the name Abies canadensis inverta in the chapter \"Plans of Residences and Grounds,\" but in the second part, under the entry on \"Hemlock Fir,\" he introduces a new name for the tree, \"Sargent's Hemlock: Abies canadensis Sargenti.\" He The author first encountered Sargent's weeping hemlock in 1970, at the Arnold Arboretum, and was instantly enamored with its unusual form. ARNOLD ARBORETUM MAP (1969); ARNOLD ARBORETUM SPECIMEN IN 1970 (10712*A)\/BOTH ARNOLD ARBORETUM ARCHIVES 12 Arnoldia 78\/2 \u2022 November 2020 goes on to describe it as being \"of an eccentric rambling nature, but well clothed in verdure,\" and he provides information about its cultivation: \"Grown without training it will probably be a broad, irregular, flat-headed tree or great bush, with an over-laying of downward growing branches like that of the Scamston elm. By grafting it well up on other trees, or by tying its leader to a stick or stake we believe it will be one of the prettiest and most picturesque of evergreens. The best effect will be produced when grafted well up on an ordinary hemlock stem.\" While Scott's use of two different names for Sargent's weeping hemlock in the longer of the two 1870 editions is confusing, the discrepancy suggests that there was a gap between the publication of the two editions. In fact, I found a review of the longer version of the book in the August 1871 issue of The Horticulturist by Henry T. Williams, which clearly suggests that the complete version of Scott's book did not come out until mid-1871. For whatever reason, this edition retained the 1870 publication date and constitutes the earliest publication of the name Abies canadensis Sargenti. In the longer of the two 1870 editions of his book, Scott also states that the plant had been \"brought into notice by H. W. Sargent, Esq., who found it growing wild on Fishkill mountain.\" I could find no reference for this specific mountain in the literature of the period, but given that the town of Fishkill lies at the base of Honness Mountain, which is shown as part of the \"Fishkill Mountains\" in period maps, it could well have been an alternative name for it. If so, then Scott is in agreement with Maxwell that Sargent's weeping hemlock was discovered on Honness Mountain. Scott and Maxwell also agree on the need to stake up Sargent's weeping hemlock in order to make it a proper \"ornamental\" plant and that without this treatment it would sprawl across the ground, eventually forming a strongly pendulous shrub. Henry Winthrop Sargent's house at Wodenethe, photographed in 1886. ARNOLD ARBORETUM ARCHIVES Sargent's Weeping Hemlock 13 One final detail in Scott's description of Sargent's weeping hemlock that should be noted appears in the appendix at the end of the second part where he lists \"Sargent's Hemlock\" as reaching ten feet tall by ten feet across under the column headed \"Usual Size 12 Years from Seed\" and thirty feet tall by forty feet across under the column \"Usual Size at Maturity.\" When I first read these numbers in the early 1980s, I couldn't figure out how Scott managed to come up with them given that they were written just two years after Hoopes published the first written description of the tree, so I chalked it up to a lucky guess. The Parsons Brothers of Flushing Scott's description of Sargent's weeping hemlock and his prescient projections about its size clearly suggest that its propagation must have been well underway in the early 1870s. Samuel B. Parsons of S. B. Parsons & Sons, Kissena Nurseries in Flushing, New York, confirmed this supposition in a lecture that he presented on November 12, 1874, to the Rural Club of New York, with many prospective clients in attendance. \"But the gem of all gems is the Weeping Hemlock,\" Parsons declared. \"If left to itself, it will remain trailing upon the ground, but if the leader is tied to a firm stake it can be carried to any reasonable height, and each tier of branches will then droop in graceful curves toward the ground.\" A year later, in October 1875, Parsons sent a letter to the editor of The Garden introducing Sargent's weeping hemlock to British audiences, using the name Abies canadensis var. pendula. Parsons's promotion of the weeping hemlock to both national and international audiences clearly suggests that he was already selling or getting ready to sell the plant to the general public. As far as I have been able to determine, however, it was the nursery owned by Parsons's brother, Robert, who first offered Sargent's weeping hemlock for sale in the fall of 1874. Some two years earlier, in the fall of 1872, the brothers had decided to split up Parsons & Sons Nursery, which they had inherited from their father and jointly operated since 1841. Samuel got half of the plant stock and established S. B. Parsons & Sons, Kissena Nurseries in a new location in Flushing while Robert took control of the other half of the stock and remained at the original nursery site but changed the name to R. B. Parsons & Co.8 In his fall 1874 catalogue, Robert Parsons listed Sargent's weeping hemlock under the heading \"Abies canadensis, weeping.\" Ten onefoot- tall plants were available for the reasonable price of eight dollars, and ten larger plants (up to two feet tall) were selling for twelve dollars. Samuel's firm, S. B. Parsons & Sons, first offered the weeping hemlock in their autumn 1877 wholesale catalogue. Both brothers clearly had a financial stake in the success of the plant and cooperated in introducing it into cultivation. One of the curiosities of the weeping hemlock history is that up until 1875\u2014after its production and sale was well underway\u2014the supposed discoverer of the plant, H. W. Sargent, had said nothing about it. He finally broke his silence in the fourteen-page supplement he wrote for the ninth edition of Downing's Treatise on the Theory and Practice of Landscape Gardening: \"Abies Canadensis pendula, or Sargenti, as sometimes called, is a very interesting and distinct variety of hemlock,\" Sargent wrote. \"It is as pendulous as a Weeping Cherry, perfectly hardy, and admirably adapted for small places, though as yet very rare, Messrs. Parsons, of Flushing, alone having plants for sale. It is a sport of our native Hemlock, found in the Fishkill Mountains.\" This brief description occurs in the supplement to the 1875 edition of Downing's book but is not included in his 1859 supplement to the sixth edition where fifty-one pages are devoted to \"The Newer Evergreen Ornamental Trees.\" This omission is significant because it suggests that Sargent did not learn about the tree until after 1859. From Fishkill to Philadelphia One of the long-standing questions surrounding the history of Sargent's weeping hemlock concerns the date when it was first propagated for commercial sale. In 1939, Arlow B. Stout of the New York Botanical Garden identified J. R Trumpy, the propagator for the Parsons & Sons Nursery, as the person who visited Fishkill and collected scions from H. W. Sargent's plant, but Stout didn't provide a date for the 14 Arnoldia 78\/2 \u2022 November 2020 Sargent's Weeping Hemlock 15 trip. Trumpy was a Swiss-born horticulturist who immigrated to America in 1856 to work for the Parsons Nursery.9 When the Parsons brothers split up the nursery in 1872, Trumpy went to work for Samuel's newly established S. B. Parsons & Sons (the name Kissena Nurseries was added later), and their very first Descriptive Catalogue, from 1873, listed him as propagator on the title page. Thanks to a recently discovered article in an 1877 issue of the Moore's Rural New-Yorker\u2014written by the magazine's \"conductor,\" Elbert S. Carmen\u2014we now know what happened when Trumpy went to Fishkill in search of Sargent's weeping hemlock: Grace is not an adjective often serviceable in descriptions of Evergreens, but it is the first that comes to mind in any attempt at describing the Weeping variety of the Hemlock spruce [Abies Canadensis pendula]. The variety is comparatively new and its history interesting. The original tree was, as we learn, in the possession of an old gentleman named BURROW. Mr. J. R. TRUMPY, the well-known propagator of one of the Parsons of Flushing, heard about it, and visited BURROW for the purpose of purchasing the tree. But BURROW would not sell. Mr. TRUMPY, thus disappointed, and having a desire to possess so promising a novelty, which only those who have their hearts in the business can understand, set out for Mr. H. W. SARGENT'S (Fishkill, N. Y.), who, he had been informed, was possessed of a small specimen which, either from a layer or graft, was derived from the original tree of Mr. BURROW'S. Mr. SARGENT was gracious to the enthusiastic TRUMPY, who left him with a pocketful of cions [sic], and from this start the Weeping Hemlock was propagated and disseminated. This stunning description of J. R. Trumpy's trip to Fishkill came as a complete shock to me and upended the traditional story of Sargent's weeping hemlock by asserting that the mysterious Mr. Burrow was in possession of the \"original tree\" and that Sargent's tree had been propagated from Burrow's plant. In light of the publicity that the tree had received prior to 1877, it's surprising that none of the earlier writers\u2014or any of those that came after\u2014 mentioned Burrow, a sign that naming him as discoverer must have been somewhat controversial. The other remarkable thing about Carmen's article is that it is accompanied by the first known illustration of Sargent's weeping hemlock, which shows a healthy young specimen grafted about five feet up on the understock. Curiously, the lingering question of when Trumpy actually visited Fishkill does not get answered until eleven years later when Carmen published a second article about Sargent's weeping hemlock, in an 1888 issue of the Rural New-Yorker, that repeated (and embellished) his earlier story about Trumpy's trip to Fishkill and described how best to use the tree in the garden.10 Carmen ended his article with \"A Note from S. B. Parsons,\" which offhandedly revealed when Trumpy's fateful visit had occurred. \"I do not know the precise age of my Weeping Hemlock, but conjecture it is 25 years old, as it was one of the first we grew when we discovered it in the grounds of Mr. H. W. Sargent in 1861,\" Parsons stated. \"My specimen is 11 feet in height and 13 feet in diameter of foliage. Its height has been obtained by training up a leader, and there is no reason why it cannot be carried 20 feet high.\" In light of this 1861 date, Scott's 1870 prediction that a mature weeping hemlock would be thirty feet high by forty feet across no longer seemed so outlandish. In addition to introducing Burrow into the weeping hemlock story and identifying Sargent's \"small specimen\" of the weeping hemlock as the source of Parsons's first propagation material, both of Carmen's articles present a negative assessment of the attempts to make the tree more ornamental by grafting it \"upon high stocks.\" His 1877 article is particularly blunt: \"But the great expectations of securing an evergreen tree-form of unique and incomparable grace, thus reasonably entertained, have not been fulfilled.\" To support this assessment, Carmen quotes Samuel Parsons as saying, \"We graft it readily upon high stock in the nursery, but it does not thrive as well\u2014the naked stem cracks and suffers and the massive foliage, like most evergreens perched on high stems, is too Facing page: Jean R. Trumpy (right) propagated Sargent's weeping hemlock on behalf of the nurserymen Samuel and Robert Parsons. Samuel (left) began promoting the plant in 1874, the same year that his brother, Robert, offered it in the fall catalogue for R. B. Parsons & Co. PARSONS (T. MEEHAN, 1887)\/ARNOLD ARBORETUM ARCHIVES; TRUMPY (AMERICAN FLORIST, 1913) AND CATALOGUE (PARSONS, R. B. & CO., 1874)\/BOTH BIODIVERSITY HERITAGE LIBRARY heavy for grace and proportion, and is beaten and tossed by the winds.\" In November of 1877, just four months after Carmen's first article came out, the botanist George Thurber published an article in the magazine he edited, American Agriculturist, which echoed Carmen's negativity about high-grafting weeping hemlocks and published the second known illustration of Sargent's weeping hemlock. The intensity of the debate about whether to graft the weeping hemlock high or low on the understock dates back to 1870 when Scott advocated grafting \"well up on an ordinary hemlock stem\" in his initial description of the tree. In 1874, Samuel Parsons implicitly supported the practice of high-grafting when he stated that such weeping hemlocks were \"more like an evergreen fountain than any tree known.\" In Carmen's 1877 article, however, Parsons came out against high-grafting, and he repeated his opinion ten years later in The Garden, an English publication edited by William Robinson. Curiously, Parsons chose to illustrate this article with an image of an extremely beautiful, twenty-five-year-old specimen growing on the grounds of his nursery that had clearly been high-grafted and trained to a stake. The fact that three prominent horticulturists expressed strong negative opinions about highgrafted weeping hemlocks suggests there must have been serious survival issues with specimens propagated this way. In addition, Parsons com- The first three illustrations of Sargent's weeping hemlock depicted specimens that had been grafted high: The first (bottom) appeared in The Rural New Yorker in 1877. The second (left) appeared in the American Agriculturist that same year. The third illustration ran in The Garden in 1887 and depicted a specimen, grafted in 1862, that was eleven feet tall by thirteen feet across. ALL BIODIVERSITY HERITAGE LIBRARY CARMEN, 1877 THURBER, 1877 PARSONS, 1887 Sargent's Weeping Hemlock 17 mented that such plants were \"too heavy for grace and proportion,\" subtly expressing his preference for the low-growing specimens that, in 1874, he had disparaged as \"trailing upon the ground.\" The first commercial sales of Sargent's weeping hemlock took place in 1874 and 1875 and were followed by the tree's first public showing at the famous 1876 Centennial Exposition in Fairmount Park, Philadelphia. In what must have been a remarkable display, 105 exhibits in the \"Ornamental Trees and Shrubs\" division were arranged in the landscape surrounding Horticultural Hall.11 In his 1878 report on the Centennial Exposition, the chairman of the Awards Committee, William Saunders, published a detailed description of eight of these exhibits, only one of which was reported to contain specimens of the weeping hemlock\u2014the Hoopes Brother & Thomas Nurseries of West Chester, Pennsylvania. Amazingly, their display featured three separate varieties of weeping hemlock: Abies Canadensis inverta, pendula, and Sargentii. Contrary to my expectations, Saunders's descriptions of both the S. B. Parsons and R. B. Parsons exhibits noted that varieties of Abies Canadensis were present but did not specifically mention any weeping types. After the exposition ended in November, the commissioners of Fairmount Park arranged to purchase the plants used in the nursery exhibits for planting in the park. According to a December 15, 1876, report by Eli K. Price, chairman of the Committee on Trees and Nurseries for the Fairmount Park Commissioners, many of the nurseries that displayed plants at Horticultural Hall\u2014including Hoopes Brother & Thomas, R. B. Parsons & Co., and S. B. Parson & Sons Co.\u2014\"were actuated by a liberal desire that their collections should remain in the Park, and offered them at prices which they esteemed little over half the cost to them. It was an object to the Commissioners to secure these permanently for our Park, to be transplanted as thinning out shall be required for their healthy growth, and they have been secured by purchase.\" Later records indicate that at least four weeping hemlocks were planted near Horticultural Hall, on a site that had formerly been occupied by the Women's Pavilion.12 Who Deserves Credit? The fact that Scott initially referred to the weeping hemlock as inverta in 1870 but quickly changed it to Sargenti suggests that there might have been an issue deciding who deserved credit for introducing the plant. This idea is supported by the story of the weeping hemlock that Carmen published in 1877, which credited the mysterious \"BURROW\" with discovering the tree. In his second article, from 1888, Carmen made a bold proposal to formalize Burrow's role over that of Sargent's: \"Now this Weeping Hemlock is catalogued as Abies Canadensis Sargentii pendula. Ought not the varietal name to be Burrowii pendula, in justice to the originator? Otherwise we should say that Mr. Trumpy's name should be given, since it was due to him rather than to Mr. Sargent that the tree was introduced.\" It took a while, but I eventually figured out who Burrow was thanks to a pair of advertisements I came across in the January and February 1875 issues of The Horticulturist and Journal of Rural Art and Taste. The advertisements\u2014for Burrow, Wood & Co., Mt. Hanas Nurseries\u2014 offered \"a few thousand grafts\" of the weeping hemlock from the \"Original Tree.\" This not only confirmed Carmen's assertion that someone named Burrow played a central role in the weeping hemlock story but also identified him as a nurseryman living in the town of Fishkill. A quick check of the 1880 census records for the town of Fishkill indicated that John G. Burrow was born in 1839 and lists his occupation as \"Hybridizer & Originator of New Variety of grapes.\" He had two partners, the brothers Isaac C. and Joseph J. Wood, both listed in the 1880 census as \"nurseryman.\"13 The Burrow, Wood & Co. advertisements raise the intriguing question of why Sargent insisted in late 1875 that the Parsons brothers were the only ones selling the weeping hemlock when he certainly must have known that Burrow, Wood & Co.\u2014located just five miles from his home in Fishkill Landing\u2014had started selling the plant earlier that year. Could it be that Sargent was annoyed that Burrow claimed to have discovered the weeping hemlock before he did and therefore chose to ignore him? This 18 Arnoldia 78\/2 \u2022 November 2020 idea is supported by two items in the advertisement: first, an unusual postscript at the end of the advertisement, \"P.S.\u2014 We were the first to send out this very desirable novelty,\" indicates that Burrow, Wood & Co. was directly challenging the Parsonses' claim to have introduced the tree into commerce; and second, by using the name pendula14 to describe the weeping hemlock\u2014as opposed to Sargenti\u2014they were rejecting proposals to attach Sargent's name to the plant. Clearly, the issue of priority had caused bad blood between Burrow and Sargent, especially in light of Carmen's 1877 statement that Burrow had provided Sargent with his first weeping hemlock. One final detail in the Burrow, Wood & Co. advertisement that should be noted is that the name of their nursery, \"Mt. Hanas,\" is an alternate spelling for what is now called Honness Mountain\u2014the same location where both Maxwell and Scott said the weeping hemlock had been discovered. An 1867 map of Dutchess County by Frederick W. Beers clearly shows \"Mount Honness Nursery, Burrow & Wood\" located about a half-mile west of the center of Fishkill. The map also shows the home of \"J. Burrow\" nestled into the south slope of Honness Mountain. I suspect that this coincidence is best explained by the fact that both Maxwell and Scott were referring to the specimen of the tree\u2014\"The Original Tree\"\u2014that Burrow had growing on his property rather than to one he had found growing in the wild. An Evolving Myth Following its commercial debut in the mid- 1870s, Sargent's weeping hemlock became something of a horticultural sensation. In 1897, fifteen years after Sargent's death, his cousin Charles Sprague Sargent, director of the Arnold Arboretum in Boston, attempted to formalize the tree's origin story in a Garden and Forest An 1875 advertisement for Burrow, Wood & Co. confirmed the role of an enigmatic character in the weeping hemlock story: John G. Burrow, a nurseryman who lived at the base of Honness Mountain. ADVERTISEMENT (BURROW, WOOD & CO., 1875)\/BIODIVERSITY HERITAGE LIBRARY; DUTCHESS COUNTY, NY, MAP (1867)\/DAVID RUMSEY MAP COLLECTION, DAVID RUMSEY MAP CENTER, STANFORD LIBRARIES Sargent's Weeping Hemlock 19 article. He noted that the plant had been found \"about forty years ago on the Fishkill Mountains, in New York, and was first cultivated and made known by Mr. H. W. Sargent \u2026 Several of these plants were originally found together and transplanted and the largest of them which I have seen is on the Howland estate, in Matteawan, New York, and is now about twenty five feet across. This variety has been propagated by grafting the branches on the ordinary Hemlock, but in a few years, the grafted plants form an erect stem and lose the dense low habit which is the charm of the original seedlings.\" Keeping in mind that Sargent's statement was written some forty years after the events described, it puts the date of the discovery at \"about\" 1857. For the first time, the article also reports that \"several plants were found together and transplanted,\" but it does not say by whom. Indeed, Sargent carefully counters Scott's 1870 suggestion that H. W. Sargent was the discoverer of the \"seedlings\" by noting that he was the one who \"first cultivated and made known\" the tree. Sargent followed his cousin's lead by not mentioning John Burrow or Honness Mountain, but he does weigh in on the high-grafting debate by expressing his preference for the lowbranched \"seedlings.\" Sargent's article is also noteworthy because it mentions that one of the original plants was growing at the Howland estate in the village of Matteawan (now Beacon), New York. This marks the first time that General Joseph Howland is mentioned in connection with the weeping hemlock, but Sargent does not credit him with its discovery. This attribution came fifteen years later, in 1912, in an unsigned article in the Arnold Arboretum's Bulletin of Popular Information written by Sargent's colleague Ernest H. Wilson:15 Many years ago, four or five plants of this form [Tsuga canadensis var. pendula] were found by Taxonomist Alfred Rehder photographed the Sargent's weeping hemlock at Holm Lea, in Brookline, in 1900. ARNOLD ARBORETUM ARCHIVES 20 Arnoldia 78\/2 \u2022 November 2020 the late Joseph Howland of Mattapan [sic], New York, on one of the mountains back of Fishkill Landing on the Hudson River and were named by him Sargent's Hemlock for his friend and neighbor Henry Winthrop Sargent. One or perhaps two of these wild plants are now living, although the variety has been much propagated by nurserymen by grafting its branches on the common Hemlock \u2026 The plant in the Arboretum on Hemlock Hill Road is a grafted plant, but at Holm Lea in Brookline there is one of General Howland's original plants. In Wilson's retelling of the weeping hemlock story, he makes several mistakes: first, he confuses Howland's hometown of Matteawan with a Boston suburb, Mattapan, and then he goes on to identify Howland as the discoverer of Sargent's weeping hemlock when no one else mentioned him in this role. The saving grace of Wilson's article is that he mentions, for the first time, that one of the original weeping hemlocks was growing at C. S. Sargent's private estate, Holm Lea. In 1923, the British horticulturist Murray Hornibrook put the finishing touches on this widely cited but factually challenged version of the weeping hemlock story in Dwarf and Slow-Growing Conifers: \"Professor Sargent informs me that the nurseryman's stock has all been produced from grafts from the four original plants found near the summit of Fishkill Mountain (near Beacon City, on the Hudson River) by General Joseph Howland about 1870. The finder grew one in his own garden at Matteawan, N.Y., gave the second to Mr. Henry Winthrop Sargent of Fishkill; the third to Mr. H. H. Hunnewell16 of Wellesley, Mass., and the fourth to Professor C. S. Sargent of Brookline, Mass. The second and third are dead, but the first and fourth have made very fine specimens.\" The Horton Hemlock Hornibrook's Sargent-approved version of the weeping hemlock story from 1923 received its first serious challenge in 1939, when Arlow B. Stout of the New York Botanical Garden announced to the world that \"the largest and presumably the oldest specimen of this type (Tsuga canadensis var. pendula) is a tree that stands in stately splendor in its original wild location on the mountainside overlooking the hamlet of Hortontown,\" about eight and a half miles as the crow flies from H. W. Sargent's home in Fishkill Landing. According to Stout, \"My first knowledge of this tree was during 1937 when it came into view as I passed by auto along the newly constructed Eastern State Parkway [now the Taconic State Parkway].\" The tree was sixteen feet tall and had a single trunk\u2014eighteen inches in diameter\u2014that was unbranched for its first five feet. Stout interviewed the owner of the tree, Joseph Horton, who told him that he had known the tree \"since sixty-five years [1874] and that it was then at least one half as large as it is now.\" In February 1980, when I first visited the Horton hemlock, it was owned by Jacob Veldhuis, who was using the tree\u2014which was over eighteen feet tall and thirty-one feet across\u2014as a kind of storage shed, a use to which it was admirably, if ignobly, suited. The pendant branches concealed no less than half a cord of wood, a hundred-gallon oil tank, a ladder, a wheelbarrow, several packages of shingles, and innumerable other artifacts of country life. The branches that formed the tree's canopy grew out from the trunk at about eight feet, and within the canopy, considerable self-grafting occurred where the branches touched one another. In his 1939 article, Stout noted that the Horton hemlock was growing \"close to a dwelling,\" but I was surprised to see that it was only about twenty feet away from the corner of the house\u2014a fact that cast some doubt in my mind on Stout's \"original wild location\" hypothesis, as did the tree's single, unbranched trunk. This doubt was reinforced by the fact that I had been told that the so-called \"Knapp house\" where the tree was growing predated the American Revolution. At the same time, however, I chose to ignore the fact that the tree was growing at the edge of a relatively steep, rocky slope where it was unlikely to have been planted. Having seen the Horton hemlock in the flesh, I felt the need to learn more about it, so Facing page: Eva Scofield, photographed in 1938 (bottom), stands with the Horton weeping hemlock. The tree grew outside of a family home that first appeared on maps as \"E. Horton, Grocery\" in 1876. The author first visited and photographed the plant in 1980. SCOFIELD\/HAMILTON, ARNOLD ARBORETUM ARCHIVES; HORTON HEMLOCK IN 1980\/PETER DEL TREDICI; PUTNAM COUNTY, NY, 1876\/PUTNAM COUNTY HISTORIAN'S OFFICE DIGITAL COLLECTION 22 Arnoldia 78\/2 \u2022 November 2020 I persuaded Jack Karnig, chief forester at the nearby (and now disbanded) Harvard Black Rock Forest in Cornwall, New York, to take core samples from the lowest branches on the tree\u2014at heights of five and six feet\u2014in order to calculate its age. The cores that Jack sent me in March of 1980 came with the following note: \"Your hemlock was a son of a b----. Twice I bored and got nothing. Finally got a reserve borer (smaller one) and managed to pull two cores.\" Under the dissecting microscope at the Arboretum, I counted 119 rings in the lower of the two cores\u2014with an average width of 0.5 millimeters\u2014which meant that the tree was at least five feet tall in 1860. In other words, the Horton hemlock was already a substantial tree when Burrow and Sargent first learned about it! While I was surprised by the 1860 date, I was still skeptical that the tree was growing in its original wild location given its single-trunk form and its proximity to the house. My suspicions were confirmed a year later when I unexpectedly discovered two photographs of the Horton hemlock in the Arnold Arboretum Archives. They were taken in May 1938 by Ormond Hamilton, a noted conifer enthusiast from Conway, Massachusetts, and the handwritten caption on the back of one of them stated that the tree was \"growing on place of Miss Eva Horton, Horton Town, Hopewell Junction, N. Y. This is not far from Beacon, N. Y. It was transplanted from mountain back of Beacon to its present site by Miss Horton's grandfather.\" I was stunned by this discovery, and in 1983, when I published my book on Sargent's weeping hemlock, I rejected Stout's theory that the Horton hemlock was the original tree in its original location and postulated instead that \"grandfather Horton discovered at least five weeping hemlock seedlings on the mountains between Hortontown and Beacon, New York. Sometime after 1859 but before 1865, he collected one plant for himself (and staked it) and sold the rest to H. W. Sargent.\" At the time, I naively thought I had finally solved the mystery of Sargent's weeping hemlock. Inspired by my book, Dennis Murphy of Warwick, New York, wrote me a letter on July 17, 1986, describing how he had visited the Horton hemlock in the company of a local dairy farmer, Vern Jackson, who told him that the house adjacent to the tree had been used as a store for many years. Murphy also spoke with Smith Townsend, one of the oldest residents in the area, who told him that Eva Horton's grandfather Alvah never lived in the house and that her father, Joseph, did not move there until \"after the death of Enoch Horton [in 1913] who was the last proprietor of the store.\" According to Townsend, Enoch, Alvah, and Joseph Horton were all buried in the cemetery located behind the old Calvary Methodist Church on Hortontown Road, and indeed, when Murphy visited the cemetery, he located the tombstones for both Alvah and Enoch. When I received Murphy's letter, I had no idea what to think given that it upended my published version of the origin of Sargent's weeping hemlock. I thanked Dennis for his letter and filed it away. And that's where things sat until 2015, when, by chance, I came across a statement by H. W. Sargent, from 1880, about who really discovered the weeping hemlock. This unexpected discovery got me thinking about the tree again and prompted me to pull out my old files where I rediscovered the letter from Dennis Murphy and the questions it had raised. One thing led to another and, with the help of the internet and several local historians, I was able to piece together the history of the Horton family farm. It turns out that the house where the tree was located\u2014now listed as 339 Hortontown Road, Hopewell Junction\u2014was not pre-Revolutionary at all but had been constructed by Enoch Horton in 1874, on an acre of land he acquired from his father, Jefferson Horton, for the price of one dollar.17 A local map from 1876 shows 339 Hortontown Road as \"E. Horton Grocery\" just as Vern Jackson had remembered. The same map, as well as one from 1854, shows Jefferson Horton's house just down the road apiece. According to Smith Townsend (as reported by Dennis Murphy), Alvah Horton lived about a half mile away from Jefferson Horton on Long Hill Road, and Alvah's son, Joseph, moved into the house on Hortontown Road after Enoch Horton's death in 1913. In 1939, Joseph Horton told Stout that he had \"known\" the weeping hemlock since 1874\u2014 when he was thirteen years old\u2014which coin- \u222b Sargent's Weeping Hemlock 23 cidentally was the date that Enoch Horton acquired land from his father and would have begun clearing the land around the weeping hemlock in order to build his house. In 1938, Joseph Horton's daughter, Eva Scofield, told Ormond Hamilton that her grandfather had transplanted the tree \"to its present site,\" but this is highly unlikely since her grandfather Alvah never lived in the house where the tree was located. Given that the Horton hemlock was at least five feet tall in 1860, the most plausible explanation for why it was growing where it was is that it had always been there. The Internet to the Rescue As stated above, my research on Sargent's weeping hemlock remained dormant until 2015, when in the course of doing internet research on the history of the introduction of Japanese plants into North America,18 I came across an article from 1880 by Samuel Parsons Jr., the son of nurseryman Samuel B. Parsons and an eminent landscape architect and horticulturist in his own right. The article was a transcript of a \"prize lecture\" Parsons delivered in Boston before a meeting of the Massachusetts Horticultural Society on January 17, 1880. In his talk, Parsons described in detail\u2014and at length\u2014 how best to use the flood of new woody plants that were coming into the market, especially Japanese species recently introduced by his father's company, Kissena Nurseries. Three-quarters of the way into his presentation, Parsons mentioned Sargent's weeping hemlock. \"If the broad-leaved hemlock [Abies Canadensis macrophylla] is somewhat stern and masculine in its outline,\" Parsons began, \"the weeping hemlock [Abies Canadensis pendula Sargentii] is essentially feminine in its graceful curves and fountain-like sprays of green.\" Parsons went on to credit H. W. Sargent for discovering the tree \"about twenty years ago [1860], near his place, at Fishkill on the Hudson, and moved by his enthusiasm and appreciation of choice ornamental trees, entrusted it for propagation to the distinguished expert, J. R. Trumpy.\" As I reached the end of Parsons's article, a paragraph appended to the conclusion of his lecture caught my attention. In it, the chairman of the Saturday morning meeting, the nurseryman William C. Strong of Brighton, Massachusetts, thanked Parsons for his lecture and then said that he was going to cut the discussion short so that the attendees could hear from \"a gentleman well known to be thoroughly versed in the subject before the meeting, and the editor of the new edition of Downing's Landscape Gardening\u2014Henry Winthrop Sargent, of Fishkill, N.Y., of whose presence he desired the Society to have the advantage.\" Strong went on to report that \"Mr. Sargent spoke first of the weeping hemlock, which was first introduced by him, and which he said was a very good 'find' by an old farmer on the mountains back of his (Mr. Sargent's) house. He has the largest tree of it, which is eight feet high, and spreads from fifteen to twenty feet. He has assisted the leader by tying it up to a stake. It is difficult of propagation.\" This brief quote\u2014a proverbial smoking gun\u2014 struck with the force of a thunderbolt. Shockingly, Sargent contradicted Parsons who, just a few minutes earlier, had claimed that Sargent had discovered the weeping hemlock. No, says Sargent, the tree was found by an \"old farmer\" who had a large specimen of it at his home. The fact that Sargent specifically says, \"He has assisted the leader by tying it up to a stake\" is undoubtedly a reference to the single-stemmed Horton hemlock. In addition, Sargent's use of the present tense indicates that the \"old farmer\" who found the weeping hemlock was still alive as of 1880. Assuming a discovery date in the late 1850s, Enoch Horton, born in 1846, would have been too young to qualify as Sargent's \"old farmer.\" But his father, Jefferson Horton (1804-1888), was still living at the time of the lecture and would have fit the bill\u2014especially given that the 1860 census lists his occupation as \"farmer.\" Taken together, all the evidence indicates that Jefferson Horton discovered Sargent's weeping hemlock growing wild on his own property. The size of the Horton hemlock in 1880\u2014 eight feet high by fifteen to twenty feet across\u2014 coupled with my tree ring data showing that the tree was at least five feet tall in 1860, strongly suggests that Frank Scott had seen the tree and used it as the basis for his prediction that Sargent's weeping hemlock would reach a mature size of thirty by forty feet. It also seems possible 24 Arnoldia 78\/2 \u2022 November 2020 that John Burrow knew about the Horton weeping hemlock and that it was the \"Original Tree\" he referred to in his advertisement from which he had produced \"a few thousand grafts.\" Hemlock Layering Around the time that I discovered H. W. Sargent's bombshell statement in 2015, I was also working on an article documenting the layering behavior of hemlocks growing wild on Wachusett Mountain, in central Massachusetts.19 My research showed that the low-hanging branches of stunted hemlocks growing on exposed, rocky sites can form adventitious roots where they come in contact with the soil and, over time, readjust their orientation from horizontal to vertical. In a moment of clarity, it dawned to me that the layering behavior of hemlocks that I had observed on Wachusett Mountain might be relevant to Jefferson Horton's discovery of the weeping hemlock. Could it be that the low-growing \"seedlings\" that C. S. Sargent first mentioned in 1897 were actually rooted branch layers dug up from the periphery of the wild weeping tree that Horton discovered? To my mind, finding a lone weeping hemlock with attached branch layers is much more plausible than finding five virtually identical mutant seedlings growing in one place. If there was just one original weeping tree sprawling across the ground, then it was probably growing on a sunny, exposed site with thin soil\u2014similar to the examples that I observed on Wachusett Mountain\u2014and its strongly pendulous lower branches would have been retained long enough to develop into layers. If this layering theory is applied to Sargent's weeping hemlock, it seems likely that when The size and age of the Horton weeping hemlock, photographed here in 1938, suggests that it was the original tree\u2014 staked in the location where Jefferson Horton found it. The photographer, Ormond Hamilton, reported that the trunk measured twenty-two inches in diameter at three feet off the ground. ARNOLD ARBORETUM ARCHIVES Sargent's Weeping Hemlock 25 Jefferson Horton discovered the weeping hemlock on a steep, rocky slope on his own property, it would have been growing prostrate along the ground. Assuming the tree behaved like the ones I saw on Wachusett Mountain, he might well have dug up a couple of the layered branches and sold them to Burrow and Sargent. He then tied a branch on the remaining plant to a stake to create a single trunk. It also seems possible that he might have induced his tree to form the additional layers by pinning its pendulous branches to the ground. Evidence for the layering of Sargent's weeping hemlock comes from multiple sources: First, many of the mature, multistemmed specimens of the tree display layered lower branches. In fact, Al Fordham, a former propagator at the Arnold Arboretum, successfully removed one such layer, in 1966, from the weeping hemlock that C. S. Sargent had planted at his Brookline estate, Holm Lea. Second, when the nurseryman Jacob C. van Heiningen20 spoke to Stout about the origins of Sargent's weeping hemlock, in 1939, he reported that he had stopped grafting the hemlock because of their poor survival rate and that he had propagated several hundred plants by \"the old fashioned way of layering which is naturally perfect, as they are on their own roots.\"21 Third, H. W. Sargent himself never used the word seedling, but instead called the plant \"a sport of our native Hemlock.\" Sport is an old-fashioned horticultural term that describes a mutant plant that obviously deviates from the normal type. In his 1874 article \"Evergreens, Novelties and Dwarfs,\" Maxwell also uses the term \"Sports of Nature\" to describe various mutant conifers and points to the weeping hemlock sport found on \"Mt. Hounes\" as an example of \"a real deformity\" that became a \"thing of beauty\" after receiving proper horticultural treatment (high-grafting and staking). Perhaps the most convincing bits of evidence for the theory that Sargent's weeping hemlock was derived from a single plant comes from the Burrow, Wood & Co. advertisement that referred to an \"Original Tree\" and from Carmen's 1877 statement that Sargent's tree at Wodenethe, \"either from a layer or graft, was derived from the original tree of Mr. BURROW'S.\" Taken together, all of these early references clearly suggest that Jefferson Horton's original discovery consisted of a single tree that he propagated by layering\u2014the \"single sport theory\"\u2014 rather than the \"multiple seedlings theory\" proposed by C. S. Sargent some forty years after Horton's initial discovery. Indeed, Sargent's statement that \"the dense low habit which is the charm of the original seedlings\" implies a level of uniformity that is more characteristic of vegetatively propagated layers than a group of genetically distinct seedlings.22 As I reported in A Giant Among the Dwarfs, there is considerable variation in the size and form of the oldest specimens of Sargent's weeping hemlock as well as considerable debate as to whether these differences are genetic or the result of horticultural practices.23 The surprisingly heated debate about the merits of highgrafting among the horticulturists of the day make it clear that the different appearances of the original specimens are a reflection of their mode of propagation\u2014layering versus grafting\u2014 and whether or not they were staked.24 The Final Story Putting all this information together, I can now present the most likely\u2014and hopefully final\u2014version of the Sargent's weeping hemlock story: Sometime in the 1850s, \"an old farmer,\" Jefferson Horton, discovered a wild weeping hemlock growing on his property in Hortontown (Hopewell Junction), New York. The tree, which was rediscovered by A. B. Stout in 1937, was growing in its original wild location about twenty feet from the house and grocery store that Jefferson Horton's son Enoch had built in 1874. Sometime prior to 1861, John Burrow learned about Horton's weeping hemlock and obtained a layer, which he planted on his own property on Honness Mountain in Fishkill. Around the same time, Henry Winthrop Sargent also learned about the weeping hemlock and obtained a layer of it from either John Burrow or Jefferson Horton. The specimens that both men were growing were relatively small when J. R. Trumpy of Parsons & Sons Nursery visited Fishkill in 1861 looking for propagation material. After Burrow refused to sell him his tree, Trumpy visited Sargent who gave him some 26 Arnoldia 78\/2 \u2022 November 2020 scions, and he grafted these when he returned to Flushing. At some point, Sargent obtained at least three additional weeping hemlock layers from either his own tree or from Horton's tree. He planted one of them at General Joseph Howland's estate, Tioronda, in Matteawan, New York; gave a second to his cousin C. S. Sargent, who planted it on his estate, Holm Lea, in Brookline, Massachusetts, in 1871; and gave the third to his kinsman Horatio Hollis Hunnewell of Wellesley, Massachusetts. Josiah Hoopes published the first description of the weeping hemlock in 1868. Frank J. Scott gave the tree its first Latin name, Abies canadensis inverta, in 1870, and later that year published the first proper description of Sargent's weeping hemlock under the name Abies canadensis Sargenti. Robert B. Parsons & Co. of Flushing, New York, was the first nursery to offer the tree for sale in the fall of 1874, and his brother, Samuel, started writing about it in horticultural magazines around the same time. Burrow, Wood & Co., Mt. Hanas Nursery of Fishkill, began offering grafts of the \"Original Tree\"\u2014Jefferson Horton's tree\u2014in January 1875 under the name Abies canadensis Pendula, the first time this name was applied to the plant. At least four specimens of Sargent's weeping hemlock were put on public display at the Centennial Exposition in Philadelphia in 1876 and were later planted out on the grounds of Fairmount Park. Elbert Carmen published the first illustration of Sargent's weeping hemlock in 1877, followed a few months later by a second one from George Thurber, and a third from S. B. Parsons in 1887. In 1937, Arlow B. Stout rediscovered Jefferson Horton's specimen of Sargent's weeping hemlock in Hortontown, about four miles southeast of Fishkill and eight miles from Beacon. The convoluted story of Sargent's weeping hemlock\u2014which should by rights be called Horton's weeping hemlock\u2014is a cautionary tale about the confusion and infighting that often surrounds the issue of who gets credit for the discovery and introduction of a new plant as well as the myth-making that sets in once the facts have been clouded by the passage of time.25 Nomenclature In 1983, I accepted Alfred Rehder's 1949 determination that the correct scientific name for Sargent's weeping hemlock was Tsuga canadensis forma pendula. I did this because of C. S. Sargent's assertion that the original discovery consisted of \"several seedlings\" found in the wild fit the technical requirements of a botanical forma.26 Because I now know that the original specimens of Sargent's weeping hemlock were actually layers from a single plant, the tree should be reclassified as a horticultural cultivar.27 In the light of this new information, the relevant question becomes what the \"correct\" cultivar name for Sargent's weeping hemlock should be rather than what rank it should be. According to Article 29.2 of the International Code of Nomenclature of Cultivated Plants,28 \"When there are two or more names in use for the same cultivar \u2026 the name that best preserves existing use is to be chosen as the accepted name by the appropriate International Registration Authority without regard to any rank in which those epithets might have been established or to the principle of priority.\" Scott's first epithet, inverta, from 1870 is clearly out of the running given that it lacked a proper description and it last appeared in print in 1876. Scott's second 1870 proposal, Sargenti, was properly described and is in wide use today as 'Sargentii'.29 Pendula came late to the party, first appearing in 1875, and seems to be used more commonly today than Sargentii. In 1983, I chose to use the name pendula because I thought that the tree was a botanical forma and the German botanist Beissner, in 1887, was the first author to describe Sargent's weeping hemlock as a forma with the name pendula. Now that I know Sargent's weeping hemlock is actually a cultivar, I prefer using the name 'Sargentii' because it helps clarify the distinction between the two categories. I also like the name 'Sargentii' because it has temporal priority and reflects the plant's common name, but it's up to the International Registrar to make the final determination. Sargent's Weeping Hemlock 27 HORTONTOWN: Based on branch core data, the single-trunked Horton hemlock was at least 5 feet tall in 1860, making this the oldest known specimen of Sargent's weeping hemlock. In 1880, H. W. Sargent said the tree was 8 feet tall by 15 to 20 feet across. In 1980, it was 18.3 feet tall by 31 feet across with a trunk diameter of 24.5 inches. When I visited the tree in December 2018, it was completely dead but still standing with a trunk diameter of 28.3 inches. A picture of the tree on the internet from spring 2015\u2014when the house at 339 Hortontown Road, Hopewell Junction, New York, was put up for sale\u2014shows it to be in poor condition. In a Google Earth image of the site on April 16, 2016, the tree appears dead. WODENETHE: Henry Winthrop Sargent purchased the twenty-two-acre parcel of land that became Wodenethe in 1841 and described the evolution of its landscape in the supplement to the sixth edition of Andrew Jackson Downing's Theory and Practice of Landscape Gardening, published in 1859. Sargent died in 1882, but the property remained in the family until 1921, when the house and grounds were sold and incorporated it into the Craig House Sanatorium. In 1955, Wodenethe was sold to a developer. The house was burned down as part of a fire-training session by the Beacon Engine Company in order to prepare the land for subdivision and housing construction. The first reference to a weeping hemlock at Wodenethe came in 1868 from Hoopes, and the last came from Maxwell, in 1874, who called it \"one of the most interesting and ornamental plants in his entire collection.\" As for the question of when Sargent's tree might have died, it is worth noting that Charles Sprague Sargent made no mention of a weeping hemlock in the article he wrote about Wodenethe in 1897. Current Status of Notable Sargent's Weeping Hemlocks TIORONDA: In 1859, Joseph Howland purchased sixty-five acres of land as a site for his country estate, Tioronda, in the village of Matteawan, on the other side of Fishkill Creek from the home of H. W. Sargent. Construction of the house was completed in 1861 while Howland was off fighting the Civil War. He returned home with the rank of brigadier general. Sargent oversaw the laying out of the grounds for Howland, and at some point, he planted a layer from the original weeping hemlock near the entrance. Howland died in 1886, and his widow sold the estate in 1911. In 1915, the property was converted into America's first privately run psychiatric center and renamed The Hortontown weeping hemlock 1981 (above) and standing dead in 2018. Note the Taconic Parkway in the background. ALL \"CURRENT STATUS\" PHOTOS BY THE AUTHOR UNLESS NOTED 28 Arnoldia 78\/2 \u2022 November 2020 Craig House.30 The facility closed its doors in 1999.31 The tree was heavily pruned in the late 1990s or early 2000s and treated for hemlock woolly adelgid (Adelges tsugae). In December 2018, the Tioronda specimen was 16 feet tall and 40 feet by 34 feet across and had four major trunks with basal diameters ranging from 16 to 29 inches. HOLM LEA: H. W. Sargent also provided a weeping hemlock to his cousin Charles Sprague Sargent, who planted the specimen at Holm Lea, in Brookline, Massachusetts. According to the caption on the back of a May 1923 photo, located in the Arnold Arboretum archives, the tree was planted in 1871. When I measured it in 1980, it was 7.5 feet tall and 32.5 feet across with multiple trunks emerging from the ground. On February 23, 1984, the tree was destroyed by a fire of suspicious origin, perhaps set by some teenagers who were reported in the vicinity of the tree that night. Indeed, the tree had long been an attraction for neighborhood children who called it \"The Fort\" and often played beneath its pendant branches. The Arnold Arboretum collected a layer off of the Holm Lea tree in 1966, and the resulting plant (accession 655-66*A) is currently 7.6 feet tall and 17.3 by 15.5 feet across with a basal trunk diameter of 16 inches. HUNNEWELL: H. W. Sargent described the making of Horatio Hollis Hunnewell's estate in Wellesley, Massachusetts, in his 1859 supplement to the sixth edition of Downing's book, in the same chapter that described the creation of Wodenethe. Hunnewell was married to Isabella Wells, H. W. Sargent's first cousin, and through this connection was also related to C. S. Sargent. Some people have suggested that a large weeping hemlock in the Hun- The Tioronda weeping hemlock in 1980 and December 2018 (top two). The Holm Lea weeping hemlock in 1980 and in 1984, with Gus Kelley, after the fire. Sargent's Weeping Hemlock 29 newell Pinetum might have been one of H. W. Sargent's original plants because of its multistemmed form, but it does not appear on an 1895 map of the collection. In 1923, Murray Hornibrook\u2014on C. S. Sargent's authority\u2014announced that one of the original seedlings went to Hunnewell but that it had died. In 2012, the estate's longtime horticulturist, David Dusenbury, uncovered a reference from the late 1920s among the unpublished writings of Theophilus D. Hatfield, who worked at the Hunnewell estate from 1887 until 1929: \"The original plant [of Sargent's weeping hemlock] I believe is still on the late professor Sargent's estate in Brookline. Our plant, of course, is a graft, and indeed a very handsome specimen, admired by all visitors.\" As of 2019, the tree measured 22 feet tall and 47.5 feet by 42.2 feet across; it has four large trunks with breast-height diameters ranging from 13 to 27 inches. FAIRMOUNT PARK: Following the end of the 1876 Centennial Exposition in Philadelphia, at least four weeping hemlocks were sold to the Fairmount Park Commission and planted near Horticultural Hall, on a site that had formerly been occupied by the Women's Pavilion. In 1896, Joseph Meehan reported that the four trees were \"a source of much interest to the numerous visitors to the park. Having been grown for twenty years, they excel [sic] probably any other specimens in these parts. They are about six feet high and eight feet in width.\" In 1939, they ranged in size from 12 to 14 feet tall. When I visited the park in 1994, all four trees were still alive, and the largest specimen measured between 34.5 feet tall and 40 by 50 feet across with a basal trunk diameter of 31 inches. In November 2018, only this tree and one other were still alive. The Hunnewell weeping hemlock in 1930 and 2010 (top two). The Fairmount Park weeping hemlocks in 1938 and one in November 2018. A. B. STOUT, ARNOLD ARBORETUM ARCHIVES H. G. MAYER, ARNOLD ARBORETUM ARCHIVES 30 Arnoldia 78\/2 \u2022 November 2020 ARNOLD ARBORETUM: A singlestemmed, grafted specimen (accession 1514-2*A) was propagated in 1881 from scions taken from a grafted plant received from S. B. Parsons & Sons, Kissena Nurseries in 1880. In 1980, a large branch with sixty-six growth rings was removed from the tree 6 feet up the stem, indicating that it was at least this tall in 1913. As of December 2018, the tree was 16 feet tall by 25 feet across with a trunk diameter at breast height of 19.4 inches; its trunk had a pronounced lean to it and structural roots near the base were protruding out from the ground. LOVE LANE: Claiming to have found the largest anything is always a risky proposition, but with that caveat, the largest weeping hemlock I have seen is growing in a lawn on a private estate in Weston, Massachusetts. It was planted in the early 1900s on property owned by John G. Freeman and his wife, Caroline Case, the sister of Marian Case, who established Hillcrest Farms at the Case Estates.32 In 1980, this giant, multistemmed specimen of Sargent's weeping hemlock was 19 feet tall and 47 feet by 43 feet wide. In 2018, it was 22 feet tall and 79 feet by 70 feet across with eight huge, ribbon-shaped stems with diameters ranging between 20 and 32 inches. It's a truly magnificent tree, but the main trunk was starting to split apart and one of its upper limbs had broken, leaving a large hole in the once closed canopy. The Arnold Arboretum's oldest weeping hemlock (1514-2*A) in September 1945 and June 2019 (top two). The Sargent's weeping hemlock on Love Lane in 2019 and, showing the branching structure, in 2016. J. F. ROCK, ARNOLD ARBORETUM ARCHIVES Sargent's Weeping Hemlock 31 DEDICATION This article is dedicated to the memory of Gus Kelley of Little Compton, Rhode Island, who first inspired me to take up the study of Sargent's weeping hemlock. Endnotes 1 Jenkins, 1946 2 According to A Book of the United States, edited by G. Mellen and published in 1838: \"The Highlands of the Hudson, or Fishkill Mountains, which first appear about forty miles from New York, are marked for their sublimity and grandeur, and interesting from their connection with many great events of the revolution. This chain is sixteen miles in width, and extends twenty miles along both sides of the Hudson.\" 3 Smith (1856) paints a vivid picture of Wodenethe in all its glory, and Spingarn (1937) documents the significant role that Sargent played in the history of American horticulture not only as a writer and plant collector but also a horticultural innovator. He was one of the first Americans to use a lawn mower and marveled, in 1855, at how it could do in eight hours what \"formerly occupied two men and a boy the better part of nine days to do, and infinitely better too.\" 4 Sargent's ideas about gardening were heavily influenced by the writings of the British horticulturist J. C. Loudon. According to Spingarn (1937), \"Loudon's 'gardenesque style' became Sargent's ideal, as it became that of the Arnold Arboretum\u2014in other words, an arboretum landscaped like a park-like English estate.\" 5 The earliest scientific name for the eastern hemlock, also known as the hemlock fir or hemlock spruce, was Pinus canadensis, bestowed by Linnaeus in 1763. Andr\u00e9 Michaux changed it to Abies canadensis in 1796, and in 1855, the French botanist L. Carri\u00e8re created the genus Tsuga to encompass all hemlocks and assigned the name Tsuga canadensis to the eastern hemlock, a change that was accepted slowly. 6 Apparently Honness Mountain is a corruption of the Dutch term hondenneus, meaning \"dog's nose.\" 7 Plans VIII, IX, XIII, XIV, XV, XVI, and XVII also feature \"Sargent's hemlock, Abies canadensis inverta.\" 8 Williams, 1872; Hoopes, 1875 9 Jean Rudolph Trumpy was born in Glarus, Switzerland, in 1830 and died on May 21, 1913; he worked in the gardens of the King of Bavaria before coming to America in 1856 (A. F. F., 1913). 10 The illustration that Carmen used with his article is of a specimen at Parsons's Nursery and first appeared in an article that S. B. Parsons wrote for The Garden in 1887; it also appeared in an unsigned 1887 article in the Horticultural Art Journal, volume 2, page 72. 11 T. Meehan, 1876 12 Rothrock, 1880; Jenkins, 1933 13 See also the 1910 obituary of Isaac C. Wood, published in Horticulture, 12(5): 156. 14 This advertisement constitutes the first use of the epithet Pendula to describe Sargent's weeping hemlock. 15 The article can be ascribed to Wilson due to the fact that he reprinted much the same information\u2014including the mistakes and much of the same phrasing\u2014in an article he wrote for The Garden Magazine in 1920. 16 Horatio Hollis Hunnewell was married to Isabella Wells, H. W. Sargent's first cousin (Sutton, 1970). 17 Sallie Sypher, deputy historian for Putnam County, located the Horton Claim Deed (executed on June 10, 1874) in Liber 67, pp. 21-22 at the Putnam County Clerk's Office. 18 Del Tredici, 2017 19 Del Tredici and Orwig, 2017 20 Van Heiningen established South Wilton Nurseries in Wilton, Connecticut, in the early 1900s. 21 See Hoopes (1868) and Wells (1955) for a description of layering in nursery practice. 22 It is tempting to speculate that the tendency of Sargent's weeping hemlock to \"come true\" from seed (first observed in 1906) provides evidence for Sargent's seedling theory (Jenkins, 1935; Stout, 1939; Del Tredici, 1983). The parsimony principle (Occam's Razor), however, suggests that propagating six layers off one parent tree is more likely than finding six identical seedlings growing in a single location. 23 Bean, 1914; Stout, 1939; Swartley, 1984 24 My own research at the Arnold Arboretum demonstrated that, after four years, grafted plants of two dwarf hemlock clones, 'Nana' and 'Cole's Prostrate', were significantly larger and broader than cutting-grown plants on their own roots (Del Tredici, 1985). Presumably these differences were due to the fact that a grafted plant is \"bi-genomic,\" with a normal root system and a dwarf top, while both the roots and the shoots of a cutting-grown plant are derived from the same dwarf genome. As regards staking, the early propagators knew that tying the leader to a stake dramatically increases both a plant's height and the speed of its growth. 25 In St. George and the Pygmies (1984), I describe the tangled story of Tsuga canadensis 'Minuta', which bears remarkable similarities to the story of Sargent's weeping hemlock. 26 According to Davis and Haywood (1965), the rank of forma (abbreviated f.) is the lowest unit of botanical classification and describes a single-character variation with a random distribution within a natural plant population. While horticultural taxonomy still uses the forma designation, it has fallen out of favor in botanical taxonomy. 27 In 1953, the horticultural concept of the cultivar was introduced as the preferred way to describe plants that have undergone some degree of human selection. Over time, the cultivar name in single quotes has largely 32 Arnoldia 78\/2 \u2022 November 2020 supplanted the use of the botanical concept of forma to describe horticultural selections. With woody plants, the cultivar name is typically, but not always, used to describe asexually propagated clones. 28 Brickell et al., 2016 29 According to the rules of nomenclature, when a plant name is derived from a person's name that ends in a consonant, the letters ii are added to it. 30 Craig House hosted many famous \"guests,\" including F. Scott Fitzgerald's wife, Zelda; Frances Seymour, the wife of Henry Fonda and mother of Jane Fonda; Rosemary Kennedy, after her catastrophic lobotomy; and the actors Jackie Gleason and Marilyn Monroe. 31 In 1933, Jenkins describes meeting Clarence Slocum, who initially managed Craig House. I met with his son Jonathan on several occasions in the 1980s, and on my last visit, he gave me the remains of H. W. Sargent's library as a donation to the Arnold Arboretum Archives. 32 According to the \"Love Lane Historical Narrative\" on the Town of Weston website, the landscape plan for the Freeman\/Paine house at 55 Love Lane was drawn up in 1901. Retrieved from https:\/\/www.weston. org\/687\/Love-Lane-Area-Historical-Narrative References A. F. F. 1913. Obituary, Jean Rudolph Trumpy. The American Florist 40(1304): 1059. See also: Anon. 1913. Obituary, J. R. Trumpy. Horticulture 17(22): 844; Anon. 1913. Noted horticulturist dead. New York Times: May 24, 1913: 4. Bean, W. J. 1914. Trees and shrubs hardy in the British Isles. London: John Murray. Beissner, L. 1887. Systematische eintheinlung der coniferen. F. L. Winterlich: Dresden. Brickell, C. D., Alexander, C., Cubby, J. J., David, J. C., Hoffman, M. H. A., Leslie, A. C., Mal\u00e9cot, V., and Jin, X. 2016. International code of nomenclature for cultivated plants (9th ed.). Leuven: International Society for Horticultural Science. Burrow, Wood & Co. 1875. Weeping hemlock grafts [Advertisement]. The Horticulturist and Journal of Rural Art and Rural Taste, 30(343): 1; and 30(344): 5. Carmen, E. S. 1877, July 21. The weeping hemlock (Abies Canadensis pendula). Moore's Rural New- Yorker, 36(3): 37. Carmen, E. S. 1888, January 21. The weeping hemlock. The Rural New-Yorker, 47: 38. David Rumsey Historical Map Collection. 1867. Map of Fishkill, Dutchess Co., N.Y. Retrieved from https:\/\/www.davidrumsey.com\/luna\/servlet\/ detail\/RUMSEY~8~1~272985~90046810:Fish kill,-Dutchess-County,-New-York?qvq=w4s:\/ where%2FDutchess%2BCounty%2B%25252 8N.Y.%252529%2F;lc:RUMSEY~8~1&mi=1& trs=50 Davis, P. H. and Haywood, V. H. 1965. Principles of angiosperm taxonomy. New York: D. Van Nostrand and Co. Del Tredici, P. 1980. Sargent's weeping hemlock reconsidered. Arnoldia, 40(5): 202-224. Del Tredici, P. 1983. A giant among the dwarfs: The mystery of Sargent's weeping hemlock. Little Compton, RI: Theophrastus. Del Tredici, P. 1984. St. George and the pygmies. Little Compton, RI: Theophrastus. Del Tredici, P. 1985. Propagation of Tsuga canadensis cultivars: Softwood vs. hardwood cuttings. Combined Proceedings of the International Plant Propagators' Society, 35: 565-569. Del Tredici, P. 2017. The introduction of Japanese plants into North America. The Botanical Review, 83(3): 215-252. Del Tredici, P. and Orwig, D. A. 2017. Layering and rejuvenation in Tsuga canadensis (Pinaceae) on Wachusett Mountain, Massachusetts. Rhodora, 119: 16-32. Downing, A. J. 1875. A treatise on the theory and practice of landscape gardening [1841] with two supplements [1859, 1875] by Henry Winthrop Sargent (9th ed.). New York: Orange Judd Co. Hoopes, J. 1868. The book of evergreens. New York: Orange Judd and Co. Hoopes, J. 1875. A visit to Parsons' Nurseries. The Horticulturist and Journal of Rural Art and Rural Taste, 30(351): 257-259. Hornibrook, M. 1923. Dwarf and slow-growing conifers. London: Country Life Ltd. Ingram, J. S. 1876. The Centennial Exposition, described and illustrated. Philadelphia: Hubbard Brothers. Jenkins, C. F. 1933. Sargent's weeping hemlock again; Hemlocks at the Centennial Exposition of 1876. The Hemlock Arboretum at \"Far Country,\" Bulletin, No.5. Jenkins, C. F. 1935. Sargent's weeping hemlock from seed. The Hemlock Arboretum at \"Far Country,\" Bulletin, No.11. Jenkins, C. F. 1946. Hemlock\u2014the queen of conifers. Arnoldia, 6(11-12): 49-60. Maxwell, T. C. 1874. Evergreens, novelties and dwarfs. The Horticulturist and Journal of Rural Art and Rural Taste, 29(337): 200-203. Meehan, J. 1896. Group of weeping hemlocks. Park and Cemetery, 5(11): 192. Meehan, T. 1876. Editorial notes: Horticulture at the Centennial. The Gardener's Monthly and Horticulturist, 18: 254-256. Sargent's Weeping Hemlock 33 Meehan, T. 1885. Editorial notes, Frank J. Scott. The Gardener's Monthly and Horticulturist, 27: 375-376. Meehan, T. 1887. Editorial notes, S. B. Parsons. The Gardener's Monthly and Horticulturist, 29: 378-379. Mellen, G. 1838. A book of the United States. Hartford: H. F. Summer and Co. Parsons, R. B. & Co. 1874. Wholesale catalogue, Fall 1874. Flushing, NY. Parsons, S. B. 1874. Evergreens. The Horticulturist and Journal of Rural Art and Rural Taste, 29(342): 362-364. Parsons, S. B. 1875. The weeping hemlock spruce. The Garden, 8: 310. Parsons, S. B. 1887, October 22. Weeping hemlock spruce. The Garden, 32: 363. Parsons, S. B., Jr. 1881. The most promising, new, hardy, ornamental trees and shrubs, and their tasteful and effective arrangement. Prize essay for the Massachusetts Horticultural Society, presented Saturday, January 17, 1880. Transactions of the Massachusetts Horticultural Society for the Year 1880: 20-43. [Reprinted in 1881 as: Rarer ornamental trees and ornamental gardens in Gardner's Monthly and Horticulturist, 23: 260 ff.] Parsons, S. B. & Sons. 1873. Descriptive catalogue of ornamental trees, flowering shrubs, fruit trees, plants and evergreens. Flushing, NY. Parsons, S. B. & Sons' Kissena Nurseries. 1877. Wholesale trade list, autumn 1877 and spring 1878. Flushing, NY. Parsons, S. B. & Sons Co., Kissena Nurseries. 1878. Wholesale trade list, autumn 1878 and spring 1879. Flushing, NY. Parsons, S. B. & Sons Co., Kissena Nurseries. 1887. Descriptive catalogue of hardy ornamental trees, flowering shrubs and vines. Flushing, NY. Price, E. K. 1876. The Michaux trees. Proceedings of the American Philosophical Society, 16(98): 340-345. Putnam County Historian's Office Digital Collection. 1876. Putnam County New York map from 1875. Retrieved from https:\/\/www.hrvh.org\/ cdm\/singleitem\/collection\/pchc\/id\/377\/rec\/2 Putnam Historical Society Digital Collection. 1854. Putnam County, NY map from 1854. Retrieved from https:\/\/www.hrvh.org\/cdm\/singleitem\/ collection\/pchc\/id\/256\/rec\/1 Rehder, A. 1949. Bibliography of cultivated trees and shrubs. Jamaica Plain, MA: Arnold Arboretum. Rothrock, J. T. 1880. Catalogue of trees and shrubs native of and introduced in the horticultural gardens adjacent to Horticultural Hall in Fairmount Park, Philadelphia. Philadelphia: s.n. Sargent, C. S. 1897. Wodenethe. Garden and Forest, 10: 449-450. Sargent, C. S. 1897. Notes on cultivated conifers, no. 11. Garden and Forest, 10: 490-491. Sargent, H. W. 1855. The Shank's lawn mower. The Horticulturist and Journal of Rural Art and Rural Taste, 10: 335. Saunders, W. 1878. Ornamental trees and shrubs. In: F. A. Walker (ed.), United States Centennial Commission, International Exhibition 1876, Reports and Awards, Group XXIX (pp. 15-29). Philadelphia: J. B. Lippincott and Co. Scott, F. J. 1870 and 1873. The art of beautifying suburban home grounds of small extent. New York: D. Appleton & Co. Smith, J. J. 1856. Visits to country places, no. 3. The Horticulturist and Journal of Rural Art and Taste, 11: 445-449. Spingarn, J. E. 1937. Henry Winthrop Sargent and the early history of landscape gardening and ornamental horticulture in Dutchess County, New York. Dutchess County Historical Society Yearbook 1937: 36-63. [Reprinted in 1938 in Landscape Architecture, 29(1): 24-39.] Stout, A. B. 1939. Weeping or pendulous hemlocks. Journal of the New York Botanical Garden, 40(475): 153-166. Sutton, S. 1970. Charles Sprague Sargent and the Arnold Arboretum. Cambridge, MA: Harvard University Press. Swartley, J. C. 1984. The cultivated hemlocks (revised by H. J. Welch). Portland: Timber Press. Thurber, G. 1877. Small evergreens for small places. American Agriculturist, 36(11): 429-430. Wells, J. S. 1955. Plant propagation practices. New York: MacMillan Co. Williams, H. T. 1871. Scott's Suburban Home Grounds. The Horticulturist and Journal of Rural Art and Rural Taste, 26(302): 238-239. Williams, H. T. 1872. Editorial notes: Parsons & Co. The Horticulturist and Journal of Rural Art and Rural Taste, 27(316): 316. Wilson, E. H. 1912, November 14. Arnold Arboretum Bulletin of Popular Information, no. 36. Wilson, E. H. 1920. The romance of our trees, no. 12: The pygmies and dwarfs. The Garden Magazine, 32(1): 36-40. Peter Del Tredici is senior research scientist emeritus at the Arnold Arboretum and the former director of living collections. The second edition of his book Wild Urban Plants of the Northeast: A Field Guide was published in March 2020."},{"has_event_date":0,"type":"arnoldia","title":"Each Year in the Forest: Autumn","article_sequence":4,"start_page":34,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25717","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24eb76d.jpg","volume":78,"issue_number":2,"year":2020,"series":null,"season":null,"authors":"Hipp, Andrew L.","article_content":"Each Year in the Forest: Autumn Andrew L. Hipp Illustrated by Rachel D. Davis HIPP, A. L. AND DAVIS, R. D. 2020. EACH YEAR IN THE FOREST: AUTUMN. ARNOLDIA, 78(2): 34-43 Bonnets I September is the attenuated tail of summer. The last flowers of great blue lobelia bloom in meadow openings or in partially shaded forest edges where they can find a little extra soil moisture. Tangles of calico aster spill into the trails, branching and short-leaved, strewn with flowers. White rattlesnakeroot flowers hang like trombone bells at the ready. Jackin- the-pulpit berries turn gradually from green to red, the masses of fruits on some plants as variable as kernels of multicolored flint corn. False Solomon's seal berries ripen from salmon to bright red and become thin-skinned and heavy with juice. Blue cohosh seeds ripen on the plant, toxic but delicious looking, a rich blue that will hold its own through winter, when you may still find an occasional seed abandoned by a gray squirrel on top of a fallen log, beside a scattering of scratched red oak acorn shells. Forest: Autumn 35 Acorn production peaks in northern Illinois around the first of the month. Nuts pile up along the trails. Many are immediately split open by squirrels or eaten by deer. Others are not eaten by mammals but are preyed upon by weevils that devour the starchy cotyledons and fill the shells with frass, exposing the baby plants to fungi and desiccation. In some cases, the only violence weevils do the seedling is to deprive it of some of the nutrients left by the mother tree in its cotyledons. Perhaps this will be enough to kill the seedling over time, leaving it too weak to hold on for a few years in the understory until there is a blowdown or an old tree falls, taking a few others with it, letting in enough light for the baby oak to photosynthesize on its own and possibly win the race to the canopy. If it fails to do so, the oak will never produce offspring of its own. The seeds falling from the tips of the tree of life in the weeks flanking the equinox are the ones we will find growing next spring. They were sparked into life in an instant of unlikely pollination. They were provisioned with food all through development. Now, we find them dispersed on feathers or fur, in the stomach of a bird, in mud lodged between toes or talons or claws. Some are dropped unceremoniously at the base of the tree to roll downhill, in a move that might appear clumsy, but what are appearances? Each species has gambled successfully over tens of thousands of generations, if not more, on that drop to the ground or that risky flight on wind, or on the passage of squirrels or jays or extinct passenger pigeons or mammoths whose interests were never identical to those of the trees. The dice drop; then the plant prepares for winter. Perhaps their seeds will germinate before they can be eaten by a vole or squirrel. If so, these notes of fall will echo for hundreds of years. II Near the end of September, the white fungal bodies of aborted entoloma sprout from the leaf litter like manna. These knots of mycelia push leaves out of the way overnight to sit on the surface of the forest floor. Over a few days, they grow into misshapen white loaves as large as an infant's fist. They are caused by a pathogenic fungus (Entoloma abortivum)1 infecting one of the honey mushrooms (Armillaria gallica).2 The latter are known best from their black, cord-like rhizomorphs that scout the soil's surface for trees to infect and then ascend the trunks beneath the bark, where they remain long after the tree is dead. Near these masses of intertwined Armillaria and Entoloma mycelia, the yellowish-brown fruiting caps of Armillaria can often be found, and sometimes the whitish caps of Entoloma as well. By night, when the rains have been just right, glowing Armillaria marks the edges of the trails, ghosts of the cambium devoured by the fungal mycelia. Rings of light mark the ends of severed boles, squeezing through passages where sunlight formerly passed from the leaves down to the roots as fixed sugars. 36 Arnoldia 78\/2 \u2022 November 2020 Stump puffballs (Apioperdon pyriforme) sprout from downed logs or form colonies in the wood chips. Then their insides turn to spores. The precocious ones desiccate and become brown inside while their peers are still white and fleshy or just turning granular inside the taut skin. Chicken of the woods (Laetiporus sulphureus) sprouts from standing dead ashes, fallen oaks, and rotting trees of several species, forming scalloped orange shelves of delicious flesh. Months hence, its bleached carcasses will mark where the fungal bodies clung bright as lanterns to the dead trunks. Chanterelles (Cantharellus sp.), bonnets (Mycena sp.), oyster mushrooms (Pleurotus sp.), and giant puffballs (Calvatia sp.) emerge and then dry or decompose over the course of a few autumn weeks. As these decomposers crowd the woods, the flowering plants become increasingly tattered. Jewelweed spanned the entire growing season, beginning as forests of nickel-sized cotyledons crowded under the leaf litter in late March and then rising into rolling hills of adult plants that dominate the landscape well into September.3 Now, it begins to yellow and wilt, thinning and breaking over. Wild ginger leaves glow with golden margins as they senesce, like autumn leaves of Ginkgo biloba. False Solomon's seal becomes variegated and stringy, the vessels running the length of its leaves draped with torn and yellowing epidermis. Sheaths surrounding the glistening black wild leek seeds split open. The seeds stare out at the coming winter for a few days before they drop to the ground. Hop sedge and Gray's sedge become decrepit, and the swollen skins of their perigynia disintegrate. Straight-stigma and curly-stigma wood sedges shatter, scattering their last achenes onto the bare soil. The white oak acorns that have made it this far lie half embedded in the soil. They split at one end, opposite the cap, cracked open by the emerging root that swells in the autumn rains. They are feeling their way blindfolded, Blue Cohosh Forest: Autumn 37 trying to get a toehold while there is still time. Their impulse to grow is strong: collect a bagful of acorns, toss it into the refrigerator next to the carrots, and keep it cool and moist and dark; even there, some will start to germinate, senselessly looking for soil. The katydids have become quiet, and the morning-time crickets purr. They and the acorns are pacing the autumn to and fro,4 getting a little work done in advance of spring. III Rain falls and temperatures fluctuate in early October. Fog pools in the prairies beneath the power lines and drapes between the spruces. The trails become sodden. Stump puffballs ripen on fallen logs or stumps, syrupy brown. Earth stars (Geastrum sp.) crank their wings out and grip the soil. Young stinkhorns (Phallus sp.) erupt, crowded together like brussels sprouts, crawling with stink bugs. Over the course of several days, they grow obscenely to several inches in height and swarm with gnats. A few days later, they become flaccid and rot. One morning last year, our woods at the Morton Arboretum were overrun by spring peepers. I started hearing their squeaks, trills, and whistles on my walk into work, their sounds shifted upslope from the wetlands where they had been calling six months earlier. There wasn't anything they could be except for spring peepers. I did not expect them, however, and I consequently could not convince myself at first that I was hearing correctly. Songbirds were migrating, and I told myself the calls I heard were those of some itinerant bird I didn't recognize. I waded into the sunflowers and towering wild lettuce to flush out any birds that might be there, but the calls stopped, as frog calls always do when you go hunting their source. They picked up again after I was safely back on the trail. After about ten minutes of this, it was clear I was hearing frogs. Peeps punctuated the woods west of Big Rock Visitor Station and all the way down to the service road that runs north through the meadow. I walked into work surrounded by them. The peepers were with us for at least a few days. Colleagues found them in leaf traps and reported hearing their songs throughout the woods at all times of the day. Chorus frogs had also rediscovered their voices and were trilling in the warm afternoons. On a cool morning midweek, I made a quick stop to listen for the spring peepers again. The forest was silent. Gnats buzzed around the stinkhorns. Then, from a hollow tucked between the shortcut trail to Big Rock and the trail that runs west along the ridge of the moraine, a single peeper called. I walked down into the hollow and poked around for five or ten minutes, but there were no other calls. In early October, sugar maple leaves are turning yellow and starting to fall. Lower branches of the American black elderberry corymbs are broken, and ray flowers fall from wingstem in the floodplains. Zig-zag goldenrod heads are pale with feathery achenes. Wood nettle leaves are chewed to lace but still have plenty of sting left. Bedraggled pale jewelweed provisions its late-season capsules, galls blistering along its leaf midveins and darkening 38 Arnoldia 78\/2 \u2022 November 2020 Calico Aster Elm-Leaved Goldenrod Gray's Sedge White Rattlesnakeroot Wild Leek Forest: Autumn 39 along one side. Wild leek has dropped about a third of its seeds. Fowl mannagrass culms are reclining. Enchanter's nightshade leaves have almost all fallen, leaving the stalks bristling with fruits. But false rue anemone, one of our iconic spring ephemerals, often begins sending up fresh shoots. The species is known to be a fall germinator,5 a rarity in our forests. Yet many people miss it in the fall,6 myself included for my first twenty-five years as a naturalist. By mid-October, white-throated sparrows pass through town on their way southward and fill the fields with \"tssts\" and whistles, marauding the shrubs for berries and insects. Near sunrise, a single bird may belt out its spring song, the bold three-toned \"Old-Sam-Peabody-Peabody-Peabody\" or the two-toned \"Oh-Canada-Canada-Canada.\"7 The territorial song sparrows join in as they are skipping town, possibly defending their territory on the way south,8 as they did on the way to their breeding grounds in the north. These discordant echoes of spring reverberate through the months of fall: frog calls, spring wildflowers emerging under the year's falling leaves, sparrows guarding territory as though it were breeding season. Signs of the changing season are deeply inscribed, paid for with the lives of individuals whose instincts weren't as well tuned. Time your emergence right, and you'll make it through winter. Time it wrong, and you may not. A million hard-earned habits comprise this business of laying up treasure on earth, where the moth and dust corrupt. These are the forest's strategies for getting through winter. Beauty is a byproduct. IV Chlorophyll molecules become unhooked from the proteins that bind them as the days shorten and the nights become colder. They become phototoxic to the leaves in which they reside. Each leaf then begins the process of autumn housekeeping, breaking the chlorophyll into harmless components that can be recycled.9 It reabsorbs nitrogen, nutrients, and basic elements that are costlier to assimilate than to recycle. As the engines of photosynthesis are disassembled and reabsorbed, carotenoids are exposed, producing the brilliant yellows of fall. Anthocyanin production picks up, producing reds and oranges that may protect the leaf from sun or insects for a few weeks.10 It is a short period of intense color, shaped by the balance of daytime and nighttime temperatures, the shortening hours of daylight, the timing of precipitation, and the internal coordination of chlorophyll degradation, redistribution of resources in the tree, and the production of new pigments. Activity at the molecular level scales up to cells, to leaves, to canopies, finally to hillsides in color. Last year, an early snow fell on Halloween, weighing tree branches down and tearing leaves off prematurely. The next morning, an hour after sunrise, yellow sugar maple leaves chirped almost inaudibly as they hit the fresh snow and glowed like lanterns on its surface. The skeletons of jewelweed were knocked to the ground. The wood nettle leaves, frozen, hung like 40 Arnoldia 78\/2 \u2022 November 2020 rags. A woodcock stopped over on its way south, skating past a twelve-inch diameter red oak that had been hauled down by the snow. Fall can be over in a moment. In most years, though, autumn funnels down to winter. Conduits between tree leaves and their branches are squeezed by a scar forming at the base of the petiole, and the trees rain resources. Leaves falling to the soil return calcium, nitrogen, and other nutrients that were shuttled upward all through the summer. Maples, basswoods, ashes, tulip trees, sassafras, and black cherries shed nutrient-rich leaves that are thin and tasty. These decompose rapidly, forming an ephemeral and semitranslucent sheet over the soil's surface. Oaks, American beeches, and shagbark hickories drop leaves that decompose more slowly, remaining on the forest floor where they insulate and provide the raw material for rodent and insect activity and the matrix for ground fires.11 The chemical composition of these leaves, particularly their calcium content, shapes the sounds we will hear the next year in the quiet evenings, as Eurasian earthworms drag whole leaves into their burrows, selecting the calcium-rich species first12 and shushing along under the leaf litter. When I stamp my foot next summer, shaking the ground, the earthworms will all slurp down into their burrows and go silent for a moment before they begin again: shh, shh, shh. June beetle grubs go dormant. Cicada children, patient by nature, gradually cease their subterranean feeding. Forest understory herbs move their resources back into their corms or bulbs or rhizomes or bequeath them to the forest floor. Bald-faced hornet queens crawl into rotting logs and prepare for winter, quiet and still. Rotten black walnut husks disintegrate in puddles at the bases of hills. Needle ice appears again in the wet soil. V By the end of November, the days are cool and overcast. White oak and red oak and sugar maple leaves interbed. A few seedlings continue twisting over soggy earthworm castings that erode to granules beneath the litter. The crickets and birds are quiet, and colors become subdued. People are mostly gone from the woods. Orion begins to show up in the evening sky, gliding upward from the eastern horizon just about the time we are settling into bed. White Oak Acorns Germinating Forest: Autumn 41 The musclewood, ironwood, beeches, and oaks rattle with marcescent leaves: the branches either mistimed or willfully ignored the last freeze of the year, and in so doing, they failed to produce the scars that would have severed these leaves from the tree.13 The squirrels have mostly gleaned the acorns and walnuts they need. Their messy nests are exposed in the treetops. Bark on many of the slender ashes and sugar maples throughout the woods is shredded where bucks have rubbed, scraping the velvet from their antlers. Jewelweed skeletons are broken over and knocked to the ground. Zig-zag and elm-leaved goldenrods are sparsely fuzzed with achenes, while white snakeroot has fully dispersed its fruits, and the few remaining bracts that once subtended the flowerheads are recoiled and twisted like starfish arms. Sandhill cranes fly southward in flocks of a hundred or more, their backs scraping the clouds. Snow comes and goes, piling up on turkeytail fungus (Trametes versicolor) and secluding itself in the bark fissures of fallen logs. Juncos and chickadees glean and then spread the persistent berries of honeysuckle and gray dogwood. They are setting next year's seedlings into motion. White avens, spinulose wood fern, hepatica, white bear sedge, and a handful of other common species photosynthesize beneath the falling snow. The rhizomes of spring wildflowers are suspended for a moment, appear to rest for winter, but extend by a hair's breadth each time the soil thaws, bending around a buried stone. The future slowly unrolls with each cell division, shaping the forest we'll walk through two and three springs hence. Leaves abscise at intervals. They gyre downward. They touch the ground. Then, there is the shush of leaves against leaves. Everything that falls accumulates and shapes the forest floor. Here, a falling tree hides the entrance to a mouse's home and crushes a mass of puffballs, and spores are dispersed. Over there, the leaves pile deeply, and then a windstorm blows them away so that the next year's fires will not burn through: as a consequence, a handful of sugar maple seedlings survives one more year in the understory. These are the endings that form the forest's beginning. Endnotes 1 Czederpiltz, D. L. L., Volk, T. J., and Burdsall, Jr., H. H. 2001. Field observations and inoculation experiments to determine the nature of the carpophoroids associated with Entoloma abortivum and Armillaria. Mycologia, 93: 841-851. And for a readable summary: Volk, T. J. 2006. Entoloma abortivum, the aborting Entoloma, a.k.a. hunter's heart, totlcoxcatl, or \"ground prunes.\" University of Wisconsin Plant Teaching Collection. Retrieved from: http:\/\/botit.botany.wisc.edu\/toms_fungi\/sep2006.html. 2 For a great article on the story of Armillaria taxonomy: Volk, T. J. 2002. The humongous fungus\u2014Ten years later. University of Wisconsin Plant Teaching Collection. Retrieved from: http:\/\/botit.botany.wisc.edu\/toms_fungi\/apr2002.html. 3 The renowned forest ecologist John T. Curtis wrote of jewelweed, \"One interesting response to light is frequently seen in mesic forests in which selective logging has been practiced so that large openings have been made in the canopy. The yellow jewelweed (Impatiens pallida) regularly forms an almost pure stand under such openings. This succulent and tender annual is very sensitive to light and is markedly reduced in height at diminished intensities. The colonies thus take on the characteristics of an integrating light meter, with the tallest plants in the center of the colony and shorter and shorter 42 Arnoldia 78\/2 \u2022 November 2020 plants toward the edges. They produce contoured mounds which reflect the chance peculiarities in shape of the canopy opening with surprising accuracy.\" Curtis, J. T. 1959. The Vegetation of Wisconsin: An Ordination of Plant Communities (pp. 122-123). Madison: University of Wisconsin Press. 4 \"Listen to the rain, more rain, treadling earth to the sodden cold wet spun heads of this room, pacing the winter to and fro.\" Borodale, S. 2012. 3rd December: Notes. Bee Journal. London: Jonathan Cape. 5 Baskin, J. M., and Baskin, C. C. 1986. Germination ecophysiology of the mesic deciduous forest herb Isopyrum biternatum. Botanical Gazette, 147: 152-155. 6 I have only dipped my toe into the woodland phenology literature, but an unpublished report by Max Partch is an interesting example. Partch took pains to observe all plant phases across numerous species well into October and still included no observations of new growth in the fall. Partch M. 1999. Plant phenology in central Minnesota. Biology Faculty Publications, 1. Retrieved from https:\/\/repository.stcloudstate.edu\/ biol_facpubs\/1 7 If you sense that the white-throated sparrow song has changed over the past decade or so, you may not be imagining it. Since 2000, the two-noted song has spread across the breeding ground in Canada to largely supplant the three-noted song, perhaps due to tutoring in the wintering grounds. Otter, K. A., Mckenna, A., LaZerte, S. E., and Ramsay, S. M. 2020. Continent-wide shifts in song dialects of white-throated sparrows. Current Biology, 30: 3231-3235.e3 8 Wingfield, J. C., and Soma, K. K. 2002. Spring and autumn territoriality in song sparrows: Same behavior, different mechanisms? Integrative and Comparative Biology, 42: 11-20. 9 Christ, B., and H\u00f6rtensteiner, S. 2014. Mechanism and significance of chlorophyll breakdown. Journal of Plant Growth Regulation, 33: 4-20. 10 The potential adaptive role of leaf coloration is an area of active study. For an informative review, see: Archetti, M., D\u00f6ring, T. F., Hagen, S. B., Hughes, N. M., Leather, S. R., Lee, D. W., Lev-Yadun, S., Manetas, Y., Ougham, H. J., and Schaberg, P. G., et al. 2009. Unravelling the evolution of autumn colours: An interdisciplinary approach. Trends in Ecology & Evolution, 24: 166-173. For a recent evaluation of competing hypotheses: Pena-Novas, I., Archetti, M. 2020. Biogeography and evidence for adaptive explanations of autumn colors. New Phytologist, 228(3): 809-813. 11 Among the species I have included here, those dominated by arbuscular mycorrhizae (e.g., the maples) tend to decompose more quickly than those by ectomycorrhizal fungi (e.g., the oaks). Phillips, R. P., Brzostek, E., and Midgley, M. G. 2013. The mycorrhizalassociated nutrient economy: A new framework for predicting carbon-nutrient couplings in temperate forests. New Phytologist, 199: 41-51. Chicken of the Woods PLANTS REFERENCED Forest: Autumn 43 Andrew Hipp is the senior scientist in plant systematics and herbarium director at the Morton Arboretum in Lisle, Illinois. He conducts research on the origins and implications of plant diversity, with a focus on oaks, sedges, phylogenetic ecology, and trait evolution. You can read about his research at http:\/\/systematics.mortonarb.org and follow his natural history blog at https:\/\/botanistsfieldnotes.com. Rachel Davis is an independent visual artist in the Chicago area. She works at the interface of natural science, abstract painting, printmaking, and textiles, integrating the formal and empirical elements of the natural world in her work. You can see more of her work at https:\/\/artbumble.com and follow her on Instagram: @art_bumble. Acer saccharum - sugar maple Ageratina altissima - white snakeroot Allium tricoccum - wild leek; A. burdickii is sometimes recognized as a distinct species, and my account also applies to that species Arisaema triphyllum - Jack-in-the-pulpit Asarum canadense - wild ginger Carex albursina - white bear sedge Carex grayi - Gray's sedge Carex lupulina - hop sedge Carex radiata - straight-stigma wood sedge (I made up this common name, because the sometimes-applied \"straight-styled wood sedge\" is a misnomer; the stigmas separate this species from C. rosea, not the styles) Carex rosea - curly-stigma wood sedge Carpinus caroliniana - musclewood Carya ovata - shagbark hickory Caulophyllum thalictroides - blue cohosh Circaea canadensis - enchanters' nightshade Cornus racemosa - gray dogwood Dryopteris carthusiana - spinulose wood fern Enemion biternatum - false rue anemone Fagus grandifolia - American beech Fraxinus sp. - ash Geum canadense - white avens Glyceria striata - fowl mannagrass Helianthus sp. - sunflowers; here the common woodland species are H. strumosus and H. decapetalus Hepatica sp. - hepatica Impatiens pallida - pale jewelweed; the description in the first half of this essay also applies to I. capensis, though I. pallida is the more common in the woods I frequent Juglans nigra - black walnut Lactuca sp. - wild lettuces Laportea canadensis - wood nettle Liriodendron tulipifera - tulip tree Lobelia siphilitica - great blue lobelia Lonicera sp. - honeysuckle Maianthemum racemosum - false Solomon's seal Nabalus albus - white rattlesnakeroot Ostrya virginiana - ironwood Prunus serotina - black cherry Quercus alba - white oak Quercus rubra - red oak Sambucus canadensis - American black elderberry Sassafras albidum - sassafras Solidago flexicaulis - zig-zag goldenrod Solidago ulmifolia - elm-leaved goldenrod Symphyotrichum lateriflorum - calico aster Tilia americana - basswood Verbesina alternifolia - wingstem 12 Holdsworth, A. R., Frelich, L. E., and Reich, P. B. 2012. Leaf litter disappearance in earthworm-invaded northern hardwood forests: Role of tree species and the chemistry and diversity of litter. Ecosystems, 15: 913-926. 13 My understanding of this phenomenon comes primarily from an unpublished University of Wisconsin-Madison botany thesis on the anatomy of marcescent leaves in black oaks. As far as I know, the only published report on the thesis is a brief article I wrote in 1996 for NewsLeaf, the newsletter of the University of Wisconsin-Madison Arboretum, then updated in 2005 as \"When Oak Leaves Fail to Fall,\" Plant Health Care Report, 2005.03: 11-12; reprinted in 2007 in the Taltree Arboretum's newsletter, Tag Along, 6: 6-7."},{"has_event_date":0,"type":"arnoldia","title":"How Trees Were Urbanized","article_sequence":5,"start_page":44,"end_page":47,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25718","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24ebb25.jpg","volume":78,"issue_number":2,"year":2020,"series":null,"season":null,"authors":"Andersen, Phyllis","article_content":"The Roman engineer Vitruvius, writing in the first century BCE, suggests that trees were the original model for columns. The classical colonnade became the precursor to the tree-lined street. The repetition of identical elements evokes a kind of control, an organizing principle for settings otherwise subject to continual change. The tree metaphor persisted into the Renaissance with the architect Leon Battista Alberti, in the fifteenth century, pointing out the similarity between the increased diameter of the bottom of columns and the root flare of the planted tree. An association of trees and streets evolved. By the late nineteenth century, tree-lined streets were part of every urban planner's tool kit. Sonja D\u00fcmpelmann examines two approaches to street tree planting in her recent book, Seeing Trees: A History of Street Trees in New York City and Berlin. In the narrowest sense, her book is a case study of two cities and their approach to trees, but in the broader context, she weaves together the overlapping perspectives of urban design, tree management, and engineering and seamlessly integrates them with shifting political and social values. Her book is not only a contribution to the history of street tree planting but an original contribution to urban history. The nature-versus-culture divide applies here as it does to much of urban landscape history. \"Cities were naturalized,\" D\u00fcmpelmann writes, \"and trees were urbanized.\" In American cities, street planting was part of the Romantic \"urban pastoral\" movement of the late nineteenth century. Advocates proselytized about bringing elements of the countryside into the city, arguing that this would offer respite from the tension inherent in city life. In New York City, trees were part of the urban sanitizing movement that created Central Park. Tree care itself still depends on the health metaphor originating in that period. Trees are evaluated in terms of health and disease. Terms like immune systems, resilience, and injury are part of tree care. Until recently arborists were called tree surgeons. Conversely, contemporary urban tree-planting practices embrace sophisticated technology to create manipulated growing conditions\u2014 an honest, transparent recognition of the unique conditions of the urban landscape. Tree species are hybridized to create selections that can withstand urban conditions. Soil mixes are created with the specificity of prescription drugs. Planting pits are engineered. It is now clear that what happens underground is as essential (if not more) to tree survival as what happens above. In American cities, grand street-tree-planting projects are still part of political campaigns\u2014 a bread-and-circus approach to garner votes in upcoming elections with no provision for aftercare. Despite lessons learned about urban planting as an ongoing process that involves nurturing young plants, providing water, and protecting trees from damage and from insects and disease, municipal governments often leave trees on their own to survive with little intervention. Advocates promote trees in terms of ecosystem services, pointing out that trees moderate local weather conditions, filter pollution, and reduce global warming. Trees symbolize civic pride and the regeneration of neighborhoods. Altogether this is a heavy burden to place on young plants. Every city has tree haters as well as tree lovers. If, on one hand, trees clean the air, on the other, they are dirty: They drop leaves and fruit on sidewalks and cars. They attract bugs. Trees block signs and Facing page: Philibert de L'Orme described trees as the original inspiration for columns in his Le Premier Tome de l'Architecture, published in 1568. SOURCE GALLICA.BNF.FR \/ BIBLIOTH\u00c8QUE NATIONALE DE FRANCE ANDERSEN, P. 2020. HOW TREES WERE URBANIZED. ARNOLDIA, 78(2): 44-47 storefronts. Although a seemingly benign activity, tree planting still attracts controversy. Tree species selected for urban streets have been transformed by research and hybridization. But the selection of tree species is still vulnerable to fads. D\u00fcmpelmann quotes landscape gardener Andrew Jackson Downing in 1847: \"There is a fashion in trees that sometimes has a sway no less rigorous than that of a Parisian modiste.\" The tree of heaven (Ailanthus altissima), once recommended as a street tree, was quickly rejected because of its overwhelming odor. Norway maples (Acer platanoides), widely planted because of their ability to thrive in stressful conditions, are now banned in some communities because of their propensity to self-seed. American elms (Ulmus americana) are lost to disease. D\u00fcmpelmann reveals how the selection of tree species is vulnerable to xenophobic reactions both in Germany and the United States. Trees are caught in the debate between native-plants-only advocates and those who champion botanical cosmopolitanism. The strength of D\u00fcmpelmann's treatment of street tree planting in New York is her ability to point out the differences between the work of municipal government, high-minded philanthropic groups, and community-based initiatives that recognize the needs of specific neighborhoods. Top-down versus bottom-up. New York's tree-planting schemes are still controlled by the New York City Commissioner's 1811 plan that overlaid a grid from Houston Street to 155th Street, ignoring the island's rolling topography. The architectural historian Hilary Ballon calls New York City's grid plan \"a living framework.\" It is the tension between the rigidity of the grid and the looseness of the crowns of trees that defines the classic New York City street. While the practical benefits of street tree planting drove municipal efforts, philanthropic groups were also aware that tree-lined streets gave the rapidly growing city a veneer of a refined environment. One of the first to join the movement was Gifford Pinchot, head of the United States Forest Service. As residents of the city, Pinchot and his wife, Cornelia, were active members of the Tree Planting Association, founded in 1897. In addition to his interest in scientific forestry, Pinchot believed that trees were \"the only form through which the residents of the city can come in daily contact with nature as we know it in the woods and fields.\" By the early twentieth century, the New York landscape had become a gendered space dominated by male professionals. D\u00fcmpelmann describes how women gained entr\u00e9e to tree-planting projects by virtue of their social position and influence. Women were valued for their roles as caregivers, child protectors, and municipal housekeepers. Tree-planting efforts The Tree Planting Association highlighted plantings on New York City's West Sixty-Eighth and West Sixty-Ninth Streets as examples of \"model tenements\" in a 1903 report. GENERAL RESEARCH DIVISION, THE NEW YORK PUBLIC LIBRARY, NYPL DIGITAL COLLECTIONS were a natural fit. Cartoonists had a field day. Later in the twentieth century, women were important leaders in groups like the Neighborhood Trees Corps and the Magnolia Tree Earth Center, which began to work in neighborhoods left behind in earlier planting efforts. African American groups, especially those in Bedford- Stuyvesant, organized local tree-planting projects to regenerate their neighborhood where the street was park space. Community groups came to resist top-down government initiatives and well-meaning but na\u00efve philanthropic efforts. Both New York and Berlin began street planting to build a healthy environment for residents. In contrast to New York City's efforts, street tree planting in Berlin is inextricably associated with destruction and loss. Berlin's important achievements in urban planning in the nineteenth and early twentieth century\u2014broad tree-lined avenues and gracious parks\u2014were destroyed by war. Trees were lost in massive numbers during World War II. Many were lost to bombing; those that remained were cut down for firewood and building materials. D\u00fcmpelmann's treatment of Berlin's rebuilding includes many small, poignant stories, from the struggle to plant trees on rubble to the protection of the city's mountain ash (Sorbus aucuparia) street trees because of the nutritional value of their fruits. The partition of the city into East and West sectors after World War II removed any possibility of comprehensive urban reforestation. It was only after reunification that renewed planting efforts could build on Germany's earlier research in scientific forestry, expanding on their admired analytic methods and fieldwork. The goal of nineteenth-century German forestry research was to increase yield, yet the basic methods of scientific analysis used for research were intriguing to tree specialists well beyond the field of forestry. German plant scientists experimented with vegetative propagation and hybridization techniques to create \"the perfect tree.\" They warned of the dangers of monoculture. Charles Sprague Sargent, the Arnold Arboretum's first director, assembled a valuable collection of German forestry manuals. Information exchange in the twentieth century between American street tree specialists and their German counterparts resulted in more sophisticated and experimental planting techniques. Ideas on tree management spanned from the individual plant to the greater tree population of a city. The American landscape architect Elbert Peets, a long-time advocate of street tree planting as an essential component of city design, collaborated with the German urban planner Werner Hegemann on American Vitruvius: An Architect's Handbook of Civic Art, published in 1922. This book provided a compendium of examples of urban forms, including the integration of trees into streets and boulevards. William Solotaroff, the New Jersey-based city forester and author of the widely distributed Shade-Trees in Towns and Cities (from 1911), often referred to German models for street planting. D\u00fcmpelmann uses the complicated story of loss and rebirth of Unter den Linden, Berlin's famous tree-lined boulevard, to mirror Berlin's fractured history. It was created in the late seventeenth century and connects the pleasure ground of the Berlin Palace to the Brandenburg Gate. Long admired as one of the great promenades of Europe, the design was referenced in Frederick Law Olmsted's 1868 proposal for the parkways of Brooklyn. Unter den Linden is now freed from the isolation of East Berlin and is being restored with its long all\u00e9e of lindens as part of the greater unification of the city. For some, there is a certain cynicism about planting street trees in cities. As D\u00fcmpelmann reflects, street trees have an \"inbuilt a priori obsolescence.\" They die. In both New York and Berlin, we see that the ability of trees to thrive is contingent on human intervention. But even given that responsibility, we no longer question that they are an essential part of urban infrastructure. We have enough confidence in urban life to no longer reference rus in urbe, the country in the city. Trees on city streets are health-enhancing; they have a strong sensory presence. But in the end, it is the power of the eye, the visual value of trees on streets that sustains their place in the city. Phyllis Andersen is a landscape historian and former director of the Institute for Cultural Landscape Studies of the Arnold Arboretum Book Review 47"},{"has_event_date":0,"type":"arnoldia","title":"A Writer's World: Fagus sylvatica 'Pendula'","article_sequence":6,"start_page":48,"end_page":49,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25719","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24e8128.jpg","volume":78,"issue_number":2,"year":2020,"series":null,"season":null,"authors":"","article_content":"An afternoon in July found me in the Arnold Arboretum landscape, on a writer's quest, looking for inspiration for new poems. I was back for the first time since mid-March, when the impact of the coronavirus became unmistakable. Masked in the brilliant summer sun, I revisited the copse of white pines atop Bussey Hill, and on my way back down the hill, I cut across the dry, dusty grass where the mansion of Benjamin and Judith Bussey (the hill's namesakes) once stood. There, I found what I had been looking for\u2014an entrance into a new world, one created by an old weeping beech (Fagus sylvatica 'Pendula', accession 22746*A). Composing in my mind, I parted the emerald curtain of branches. Inside was a space of light and awe. Sequins of sun edged through a jangle of leafy streamers. At my feet, swollen roots appeared to be burnished like antique pewter. The tree forms a living memoir, written in the layering of branches that produce younger trunks. Those offspring encircle the mother trunk and echo its smooth gray. This was truly a tree to write about, with an allure both glorious and otherworldly. For me, all beeches have an aura of magic, but this tree, with its resplendent sanctuary, is my delight. It draws me in, hinting of a mythical forested world. Artists paint beeches; writers write about them\u2014and also on them. Their wide boles of smooth silver have beckoned lovers and poets through the centuries. In Shakespeare's As You Like It, Orlando hangs his love notes upon the trees, amorously declaring, \"O Rosalind, these trees shall be my books.\" This weeping beech is a well-annotated tree, incised with the names, initials, words of those who hoped to leave some mark, proclaim passion, or silently (!) voice an observation. My own words would never find a \"voice\" on a tree. Still, I am curious about the R's and E's, hearts, and watchful eyes on this trunk\u2014and I wonder about the impassioned sentiments that have already elongated and faded into its skin. When the tree was first mentioned in the Arboretum records, in 1942, it was described as \"an old tree,\" presumably part of the nineteenthcentury landscaping. The Bussey mansion was transferred to Harvard from the family in 1896, after the death of Thomas Motley, the husband of the Busseys' granddaughter, Maria. From that time until ours, how many must have marveled beneath this canopy? Weeping beeches have long inspired writers to mold that marvel into words. Garden and catalogue writers of the nineteenth and early twentieth century featured the weeping beech frequently, embellishing its description with curious and sometimes contradictory adjectives. Consider Albany Nurseries' 1915 description: \"quite ungainly in appearance \u2026 of wonderful grace and beauty.\" One wonders that they sold. Frank J. Scott, in The Art of Beautifying Suburban Home Grounds of Small Extent, published in 1873, resolved the contradictions into an enlightened use of prose: \"It is the very embodiment of all the odd freaks of growth that make trees picturesque, and the vigorous healthfulness of foliage that makes them beautiful.\" An etching in The Gardeners' Chronicle, from 1870, catches my own writing imagination\u2014 the tree leans and agitates, even in the stillness of an illustration. Its branches, from the very top to the thick undulating midsection, appear to swoop and splay about the ground in a hoary tapestry of leaf and limb. The tree's form and aspect appear as a landscape upon a landscape\u2014so yes, as the accompanying article proclaims, \"both grotesque and picturesque.\" Our tree stands steeple-like on the hill, catching a mosaic of sun. This specimen is surely, to echo a description from The Horticulturist in August 1872, \"like a cathedral built by one of the old masters of architecture.\" I consider the wonder of its life. It reassures me, even in our present world, that we, with this beech, remain, survive, hold to our roots. Fagus sylvatica 'Pendula' is evidence of nature's endurance and humanity's desire to be remembered. It is a witness. Though it does weep, I believe it is with a wondrous joy where it touches the earth. Sheryl L. White is coordinator of visitor engagement and exhibitions at the Arnold Arboretum. Her poetry chapbook, Sky gone, was published by Finishing Line Press this fall. A Writer's World: Fagus sylvatica 'Pendula' Sheryl L. White"},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25696","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270a725.jpg","title":"2020-78-2","volume":78,"issue_number":2,"year":2020,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"New Life for Old Collections","article_sequence":1,"start_page":2,"end_page":5,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25708","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d2708925.jpg","volume":78,"issue_number":1,"year":2020,"series":null,"season":null,"authors":"Clement, Wendy L.","article_content":"Facing page: Six of ten violet specimens collected by Nelson Pepper in the spring of 1892. Clockwise from upper left: Viola tricolor, V. blanda, V. pubescens, V. sagittata, V. pedata, and V. rostrata. HERBARIUM OF THE COLLEGE OF NEW JERSEY (IMAGES CROPPED) When I first joined the faculty at The College of New Jersey, in 2012, I knew we had a small collection of herbarium specimens available for use in our classes. The specimens were tucked away in the cabinets of the botany lab. At the time, my attention was turned to setting up my lab and continuing my existing plant systematics research agenda, but five years later, a planned renovation of the area where the specimens were kept gave me a reason to sort through these collections more carefully. Upon doing so, my students and I discovered a fascinating story about the origins of this small collection, a more than century-old link to botanical studies in the area, and a reminder of the value of preserving historical specimens that document how plant life is responding to a changing world. Within the collection were nearly 450 specimens collected during the late nineteenth century by three students enrolled in what was then the New Jersey State Normal School. One of the students, Nelson H. Pepper, had collected more than one hundred specimens in the spring and summer of 1892. Some of his specimens were exhibited the very next year at the 1893 Chicago World's Fair, bringing examples of the plants of Trenton, New Jersey, to an international audience. As I pulled out one of the first specimens, I noticed that the unmistakable pink anthers of the spring beauty (Claytonia virginica) were still visible under the white petals of the flower. The paper, thinning and stained with a shadow of the plant pressed upon it, could no longer support the weight of this 115-year-old specimen. This page was one of a hundred or more specimens that were kept together by a tattered leather binding. Nelson Pepper's name was still legible in gold print on the spine, along with the word \"HERBARIUM.\" Several folders were loosely stacked in alphabetical order inside the leather cover, each marked with botanical New Life for Old Collections Wendy L. Clement family names. The violet family (Violaceae) was located toward the bottom of the pile, with ten specimens of flowering violets collected in late April and May of 1892. Each plant was pressed and arranged carefully, permitting this set of specimens to illustrate the major differences of Viola species he collected in New Jersey. Each heart-shaped leaf of Viola striata and each arrow-shaped leaf of V. sagittata were separated to show the entire outline of each leaf, and the highly dissected leaves of V. pedata were similarly spread across the page. Nectar spurs\u2014 short and rounded in V. cucullata and long and pointed in V. rostrata\u2014extended out the back of the bilaterally symmetric flowers. The two other books from the same era, compiled by Sarah Elizabeth Kandle and Margaret Todd, also included over one hundred specimens. Our understanding is that all three collectors were completing an assignment for their botany class, led by professor Austin Apgar. In Apgar's more than forty-year career at the college, he was the botany and zoology instructor and, later in his tenure, the vice principal. All the while, he was a strong advocate for the establishment of the New Jersey State Museum. In his book Trees of the Northern United States: Their Study, Description and Determination for the Use of Schools and Private Students, published in 1892, Apgar presented a text for educating specialists and nonspecialists alike in botany. In the opening pages, he alludes to his own pedagogical approach of immersing students in studying botany and natural history. \"Teach [the student] to employ his own senses in the investigation of natural objects, and to use his own powers of language in their description,\" Apgar writes. Standing in his place a century later, with twenty-first-century students in my research group, I have continued the tradition of asking students to observe and document various aspects of plant diversity, such as 4 Arnoldia 78\/1 \u2022 August 2020 floral morphology or geographical distribution, as they begin to explore possible directions for their own work. After subjecting our collection to a typical freezing regimen applied to any specimen that has left an herbarium (to eliminate mold or insects), we relocated these books to archival boxes in the single herbarium cabinet in my lab. We then began the careful process of recording information from these specimens, wearing gloves to prevent further damage to the paper. The collection was rather typical of older herbarium specimens, having handwritten labels with little more information than a plant name, collection date, and vague locality. Herbarium labels now regularly include more information documented by the collector, such as detailed descriptions of the plant at the time of collection, robust accounts of the locality and habitat of the plant, and references to co-occurring species. Collectors increasingly include GPS coordinates, especially now that these data are collected with a tap on the screen of a smartphone. Most herbaria are likely to have more-recent, higher-quality collections of the nearly three hundred species documented in our herbarium books. Yet, these historic specimens represent an important snapshot in the history of the landscape surrounding Trenton, which has undergone significant changes over the past century. The specimens offer unique data points for the occurrence of these species and the developmental stage of the plant at the time of collection. My undergraduate research team often uses herbarium specimens in their work. The College of New Jersey is a primarily undergraduate institution, and at any given time, my lab is comprised of six or seven undergraduates engaged in multi-semester research projects directly related to my ongoing studies in plant systematics. Students working with me begin by engaging with projects that match their interests and then take their investigations in new directions often inspired by their own observations from herbaria or living collections. We are fortunate that our college is located near major herbaria such as the New York Botanical Garden's Steere Herbarium and the herbarium of the Academy of Natural Sciences of Drexel University, which have holdings of 7.8 and 1.4 million specimens, respectively. And now, as a result of major efforts to digitize museum collections, my students can access images of specimens from herbaria across the globe while sitting in the lab. Yet, as my students embarked on extracting data from the historic sheets, deciphering the handwritten labels and updating names to reflect the latest taxonomic changes, this time they had a direct connection to the collectors. In reflecting on their experience transcribing data from these specimens, student collaborators Linda Zhang and Aaron Lee wrote, \"Between the faded illegible cursive and yellowed paper we got a glimpse of the lives of collectors and students that spent their time gathering, identifying, and preserving these records with little knowledge that they would be stumbled upon over a century later.\" A third student, Matthew Fertakos, came to see this collection as a way to think about how individuals of the same species, divided by time, may change their biology in response to the environment. Matthew had become fascinated with published studies that used herbarium specimens to document how important phases of a plant's life cycle, such as flowering, may have changed over the past century in conjunction with changes in climate. As a DaRin Butz Intern at the Arnold Arboretum, he learned to generate maps that show the predicted distribution of a species based on locality data gathered from herbarium specimens and corresponding climate data for the year the plant was collected. Combining these two interests, Matthew asked what changes were happening in the rare but notable ecosystems of his home state, New Jersey, such as in the Pine Barrens. To date, his work has incorporated over eighteen hundred herbarium specimens, some dating back to the same era as our small collection. Many of the specimens were obtained from the Chrysler Herbarium at Rutgers University, the same institution that generously assisted with digitizing our own small collection. Focusing on a dozen herbaceous species native to the Pine Barrens, Matthew has used these herbarium specimens to generate distribution maps and estimate the first flowering date for many years over the past century. His work continues to test for correlations between changes in first flowering dates and shifts in climate to understand why some species native to the Pine Barrens now exhibit earlier fl owering dates than the century prior. Matthew's work, inspired by this small collection, joins an ongoing movement that demonstrates the hidden potential of these historical artifacts to provide information about the effects of climate change on a plant's biology. Over the past decade, herbaria worldwide have prioritized efforts to digitize their collections, increasing accessibility not only to botanists but to all scientists whose work could benefi t from these data. The renewed life that digitization has brought to museum collections has also allowed us to establish our own herbarium at The College of New Jersey, registered with Index Herbariorum, the international registry for herbaria. Now, our collection, currently focused on historical plants of the Trenton area, will soon be accessible to all. Within this digital collective, our small collection is more powerful than it could have been alone. The fact that our collection was preserved, waiting for students to use them, was not a coincidence. The families of collectors saw value in these specimens. Rather than ignoring or discarding the specimens among other attic keepsakes, the families donated these otherwise dusty old books of plant pressings back to our department. Now, more than a century after the specimens were mounted and nearly a quarter-century after the collections returned to our department, we are in a unique position to be able to breathe new life into these plants and use them in our quest to understand the effects of climate change on biodiversity, assessing changes that have happened over the past and predicting changes that will happen in the future. The story of how the collections returned to us is a testament to the value and power of amateur botanizing (what we would today call \"citizen science\") and experiential fi eldwork as part of an undergraduate education. As an instructor, I share Apgar's emphasis on engaging students in hands-on observation and documentation of the natural world, and I stress to students the importance of preserving these botanical legacies. Now, the historical plant collections from our region of the country will join the millions of specimens available digitally as the botanical community continues to ask more questions about biodiversity in an ever-changing landscape. Wendy Clement is an associate professor of biology at The College of New Jersey. She is a plant systematist and evolutionary biologist, and her current research focuses on the evolution of fusion in honeysuckles. She is a current James R. Jewett Prize awardee at the Arnold Arboretum. Two violet specimens collected by Sarah Elizabeth Kandle in 1894: HERBARIUM OF THE COLLEGE OF NEW JERSEY (IMAGES CROPPED) Viola pedatifi da (left) and V. pubescens."},{"has_event_date":0,"type":"arnoldia","title":"Model Maples","article_sequence":2,"start_page":6,"end_page":15,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25709","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24ea328.jpg","volume":78,"issue_number":1,"year":2020,"series":null,"season":null,"authors":"Grossman, Jake J.","article_content":"For all of human history and many millions of years before it began, the forests of the temperate Northern Hemisphere have been populated by maples. Today, the maple genus (Acer) extends its reach from Guatemala to Canada, the Mediterranean to Scandinavia, and Southeast Asia to the Amur Valley. Like oaks, willows, and birches, among many other genera, the maples as we know them today differentiated from their nearest relatives at a time when the global climate was hotter and wetter than today's and have since survived a long period of cooling and drying, including many ice ages. Their evolution as a genus occurred through geographical radiation across the Northern Hemisphere, interspersed by extinctions and range retractions when climatic conditions became inhospitable. Contemporary maple diversity is the result of this history and represents only a single, still snapshot from a larger, unspooling reel. The extant maples have adapted, by and large, to climatically temperate conditions: warm summers and cold winters, with occasional dry periods interspersed with regular precipitation. But contemporary, human-caused climate change is rapidly reconfiguring this climate to a warmer one with less regular and more extreme events of rain or snow, making freakish droughts, early arrivals of spring, and warm winters more common. As the climate changes, maples, like other forest species adapted to the temperate north, face an uncertain future. In my research at the Arnold Arboretum, I make use of publicly available data, existing scholarship, and, most importantly, the Arboretum's collection of over six hundred maple trees (which is nationally accredited by the Plant Collections Network) to predict how the genus will respond to climate change. Specifically, I ask how maple species differ in their response to dry soil conditions and to the shorter, warmer winters that will likely become typical in the Northern Hemisphere. In doing so, I treat maples as a model for other kinds of temperate trees. The genus makes for a good model for several reasons. First, the maples are highly diverse. The genus consists of 120 to 160 species depending on the taxonomic authority, with half of these growing at the Arboretum. Second, maples have a very wide geographic distribution, unlike some temperate genera confined to only certain continents or regions. And third, as ecologically foundational, long-lived trees, maples are of interest in and of themselves, and so there is an existing body of research addressing their natural history, ecology, and evolution. Yet, in the final analysis, the story of the maples is powerful because it is typical: the genus is neither wildly more nor less vulnerable to climate change than other temperate woody genera. As such, maples can serve as a bellwether for other temperate trees: where goes the maple, so go other temperate taxa. Thus, the genus can tell us about the past, and potentially the future, of northern forests. Paleogene Origins Maples belong to the highly diverse angiosperms, or flowering plants, which probably had differentiated from their ancestors by the beginning of the Cretaceous period, some 145 to 66 million years ago (Coiro et al., 2019). During that period, the earliest flowering plants spread across the globe, competing with and living alongside the previously dominant woody gymnosperms (including pines, cypresses, and ginkgoes). But it was during the next geologic period, the Paleogene (66 to 23 million years ago), that maples split off from their relatives among the flowering plants and truly came into their own. By the beginning of the Paleogene, the world's continents were more or less in their present locations, although their climates and Model Maples Jake J. Grossman GROSSMAN, J. J. 2020. MODEL MAPLES. ARNOLDIA, 78(1): 6-15 Facing page: Historic maple diversity has arisen over the past sixty million years\u2014a period of extreme climate fluctuations. The physiological adaptations of modern-day maples are therefore a record of that history. The redvein maple (Acer rufinerve), pictured here, is native to Japan. PHOTO BY WILLIAN (NED) FRIEDMAN 8 Arnoldia 78\/1 \u2022 August 2020 the degree of connectivity among them differed from conditions in the present day. Land bridges between North America and both western Europe (through Greenland) and East Asia (through Alaska) emerged periodically during cooler parts of this climatic cycle and sank back beneath the waves during warmer ones. In general, the global climate was hot and wet: 18\u00b0F (10\u00b0C) hotter on average than global temperatures during the twentieth century. This means that tropical biomes extended across much of the Earth's land surface, with the poles experiencing temperate conditions like those we now have at the midlatitudes. Ice was absent\u2014or very scarce\u2014on the Earth's surface. As a result, the growth and abundance of plants living at the North and South Poles was probably limited not by cold temperatures but by the scarcity of light (Tiffney and Manchester, 2001). In these conditions, so different from those we experience sixty million years later, the population of trees that would give rise to the modern maples became distinct from its kin. Per fossil evidence\u2014the appearance of recognizably maple-ish leaves and fruits\u2014and complementary modeling based on the genetics of existing maple species, it was at this point that maples diverged from other genera in the soapberry family (Sapindaceae). This group, which also encompasses horsechestnuts and buckeyes (Aesculus) and lychee (Litchi chinensis), presently consists of over 130 genera and close to two thousand species. Of these, the maple genus is most closely related to Aesculus and to Dipteronia, the two extant species of which can be found in mainland China. Indeed, China is likely the evolutionary cradle of maples; despite some fossil evidence that maples originated in North America and spread to Asia over Pacific land bridges, the most recent molecular evidence points to an Asian origin (Li et al., 2019). From these beginnings in China, maples radiated across the entire Northern Hemisphere while the warm, wet climate of the Paleogene was at its acme. Studies of fossil evidence marshaled by paleontologists such as Toshimasa Tanai (1983), Jack Wolfe (Wolfe and Tanai, 1987), and Harald Walther (Walther and Zastawniak, 2005) indicate that, during this time, the maples were highly diverse and cosmopolitan in their distribution. For example, the maple flora of western North America, for which the fossil record is particularly strong, currently consists of three species: bigleaf maple (Acer macrophyl- Maples have a widespread distribution throughout the temperate Northern Hemisphere. This map was prepared for The Red List of Maples (2009). BOTANIC GARDENS CONSERVATION INTERNATIONAL GLOBAL MAPLE DISTRIBUTION Model Maples 9 lum), vine maple (A. circinatum), and Douglas maple (A. glabrum). If generous, we could also include in this count the widespread box elder (A. negundo) and the western bigtooth maple (A. grandidentatum), which is often, and rightly, I would argue, treated as a subspecies of sugar maple (A. saccharum). Regardless, Wolfe and Tanai (1987) report paleontological evidence of ninety-one distinct maple species in the region; some of these may be the ancestors of the modern western maples, but the vast majority have been lost to extinction. This pattern, in which current maple biodiversity represents a small subsample of a formerly diverse flora, is perhaps best documented in western North America, but it likely holds true across the maples' distribution. But why? Maples on Ice In short, maples can best be thought of as either pitiable victims or, perhaps, resilient survivors of tens of millions of years of adverse climate change. Starting roughly fifty million years ago, the Earth entered a long period of gradual and intermittent global cooling, one we would still be in if not for anthropogenic climate warming. During this time, the poles and middle latitudes became cooler and drier, giving rise to the ecosystems that we now associate with the high latitudes. Permanent ice formed in the Arctic, and glaciers periodically developed and spread south. As a result, maples were pushed toward the equator in some cases and restricted to small refugia\u2014areas of permissive warm and wet conditions\u2014in others. Those species that could not tolerate the increasingly cold and arid climate or migrate away from local, harsh conditions went extinct. At the same time, ice formation and climatic cooling opened up new land bridges. These included not only those among continents but also smaller regional bridges, connecting, for instance, mainland China to Japan, Taiwan, and the bulk of the Korean Peninsula. During these moments of connection, the maples' migration in response to climate change occurred alongside the interchange of previously isolated floras. Yet the decline of global maple diversity with climatic cooling and drying was not uniform. In general, the last fifty million years have been easier on the East Asian maple flora, which, protected by the geographic diversity and relatively stable climate of the region, now includes the native range of upwards of 80 percent of today's maple species diversity. The maple floras of Europe and North America, on the other hand, have been much more vulnerable to climatic cooling, which has frequently led to considerable glaciation of both continents. However, it is important to note that these cold, dry periods of migration, extinction, and exchange were likely cyclic. As a result, maples, like many other temperate tree lineages, were squeezed and pushed, but then given periods of ten million years or so of relaxed, permissive climatic conditions. During these relaxed periods, populations likely rebounded, beneficial climatic adaptations spread, and species were able to expand from their refugia and southern havens to repopulate the north. We can see evidence of this pattern if we consider the most recent glacial cycle, alternately referred to as the Pleistocene Ice Age or the Last Glacial Period, which ended eleven thousand years ago. At the height of this Ice Age, glaciers reached well into the northern United States, and much of what we now think of as forestland was probably devoid of tree cover. During this time, sugar maples and box elders migrated deep into Central America, returning north as the glaciers retreated and the climate warmed, rendering their southern refugia too hot and dry and opening up new territory in what is today the United States. As a result, relictual pockets of these maple species can still be found in cool, wet locations, such as cloud forests, in Mexico and Guatemala. This pattern of range shifts and adaptation to new conditions serves as a likely illustration of other maple species' responses to climate change over the last fifty million years. Classification from Evolution Against this backdrop of global change and migration, sixty million years of evolution has given rise to our current maple flora of roughly 120 species. (I prefer to stick to a relatively low estimate of maple species diversity. Higher species counts\u2014close to 160 in some cases\u2014 treat two maple populations separated by geography but capable of interbreeding as different 10 Arnoldia 78\/1 \u2022 August 2020 species instead of subspecies.) As noted above, the majority of maples (more than one hundred species) are native to the genus's ancestral East Asian home; nine are native to North America; and eleven are native to Europe and Western Asia. Furthermore, a handful of East Asian species are truly tropical, extending into mainland Southeast Asia and Indonesia. Maples are a staple of the Northern Hemisphere's temperate forests, although their ecological role varies from canopy-spanning dominants (sugar maple, Acer saccharum, and red maple, A. rubrum, in the eastern United States) to specialists that are more sparsely distributed in the understory (moosewood, A. pensylvanicum) or generally riparian (silver maple, A. saccharinum, and box elder, A. negundo). Western botanists since Linnaeus have studied this considerable diversity among the maples (de Jong, 1994). Yet recent advances have finally made it possible to describe the genus in properly evolutionary terms. For many contemporary biologists, one of the main goals of taxonomy\u2014the classification of organisms\u2014should be the creation of a system in which species are organized according to their evolutionary relationships. In such a phylogenetic approach, species in a given genus, for instance, are all descended from a common ancestral population and are thus more closely related to each other than to other species outside of the genus. This is almost certainly the case for Acer as it has been described since the authoritative taxonomy by German botanist Ferdinand Pax in 1885. His work, of course, was carried out shortly after Charles Darwin's proposal of adaptive evolution and many decades before the advent of modern genetics, and so is based entirely on morphological comparisons. Since Pax, students of the maple genus have continuously refined the organization of Acer, proposing and dismissing a variety of schemes in which the genus is organized into sections (each containing species more closely related to each other than to those in other sections) and, within sections, series. For instance, since 1933, botanists have generally agreed that the morphologically similar red and silver maple, both native to North America and unique in their flowering phenology, are members of a distinct section, Rubra (de Jong, 1994), with only a single, long-lost East Asian cousin (A. pycnanthum). More recently, such classifications have been put to the test through the application of modern genomic analyses. Most recently, botanist and former Arnold Arboretum senior researcher Jianhua Li, presently at Hope College, has capped off two decades of research into maple systematics by publishing, with colleagues, a definitive phylogeny of the genus (2019). Their portrait of the genus's diversity suggests the existence of sixteen sections, most of which had become evolutionarily distinct by roughly thirty-three million years ago. This point, marking the transition from the Eocene epoch (which began fifty-six million years ago) to the Oligocene epoch (which ended twenty-three million years ago), also coincided with a dramatic drop in global temperatures following a gradual cooling during the Eocene. By the time global cooling preceding our current age had really begun to accelerate, the maple genus had experienced its most profound evolutionary diversification. The emergence, over the subsequent thirty million years, of today's maple species, was likely shaped by smaller-scale adaptations and the extinction of existing lineages, rather than by wholesale innovations within the clade. So today, after tens of millions of years of evolution, what visible traits define a maple tree? Leaf arrangement and shape, and seed type are probably the best way to identify a member of the genus. To begin with, all of the temperate maples are deciduous and broad-leaved, with opposite leaves setting them apart from many other angiosperm taxa. Most have simple, palmately veined leaves with anywhere from three to thirteen lobes, giving rise to the \"maple leaf\" shape popularized by the Canadian flag Facing page, clockwise from upper left: Leaf and flower characteristics of maples are quite variable. The clade (section Pentaphylla) with three-flowered maple (Acer triflorum) diverged from section Acer, which includes the morphologically distinct sugar maple (A. saccharum), about 29 million years ago. These groups, in turn, separated from section Rubra, which contains the red maple (A. rubrum), about 34 million years ago. Although their foliage and flowers are easy to differentiate, horned maple (A. diabolicum) and hornbeam maple (A. carpinifolium) are in two allied clades. Vine maple (A. cissifolium) has leaves that resemble the three-flowered maple, but the vine maple's clade (section Negundo) is quite distinct, having separated from its look-alikes about 63 million years ago (Li et al., 2019). Acer triflorum Acer saccharum Acer rubrum Acer carpinifolium Acer diabolicum Acer cissifolium JONATHAN DAMERY KYLE PORT DANNY SCHISSLER DANNY SCHISSLER SUZANNE MROZAK WILLIAM (NED) FRIEDMAN 12 Arnoldia 78\/1 \u2022 August 2020 and currency, which portray the leaf of a sugar maple\u2014and, in some cases, erroneously, that of an invasive Norway maple (Acer platanoides). Yet not all maple leaves fit this rubric. A few species, such as the hornbeam maple (A. carpinifolium) bear simple leaves with pinnate venation. And two surviving lineages have developed compound leaves. These species include the North American box elder in one lineage and Arboretum classics such as the East Asian paperbark (A. griseum) and Nikko (A. maximowiczianum) maples in the other. Furthermore, all maples produce beloved paired samaras: dry, winged fruit that can \"helicopter\" away from their mother tree when ripe. The presence of a paired samara generally will mark a temperate tree as a maple, though other genera, including ashes (Fraxinus), produce unpaired samaras. A tree bearing opposite leaves and paired samaras, then, is very likely to be a maple. On the other hand, flowers and bark are so diverse within the maple genus as to be unhelpful to most casual plant taxonomists. All maples produce regular, five-part (or rarely four-part) flowers, with fertilized female flowers eventually giving rise to samaras. Yet, here the similarities end. Flowers can be red, yellow, or green, and male or female (though male flowers often bear undeveloped ovaries). Maples can be dioecious\u2014single-sexed\u2014or monecious\u2014 having male and female flowers on the same plant. Monecious trees can produce waves of flowers over a single season, going from male, to female, to male again. Few trees, most notably box elder and the horned maple (Acer diabolicum), are fully dioecious, meaning they consistently present as either male or female. Maple bark presents another lesson in the genus's surprising diversity. The scaly motley of green and brown lining the trunk of European sycamore maple (Acer pseudoplatanus) makes these trees easy to spot, although in some cases, misleadingly similar to true sycamores (Platanus). The bright orange, peeling bark of the paperbark maple (A. griseum) is one of its outstanding merits as a horticultural tree. But it is the species of the section Macrantha, the snakebark maples, whose green, smooth-to-furrowed bark is, to me, most unusual and appealing. These species are restricted in their distribution to East Asia except for the North American moosewood maple (A. pensylvanicum). Hot and Cold Despite long-standing celebration of maples' morphological diversity, ecologists still lack a clear understanding of physiological diversity in the genus. I am interested in this question out of the need to forecast how particular species of maples, as well as other temperate trees, will respond to climate change. Recent studies have already documented some climate-related shifts in maple distributions. For instance, in North America, red maples seem to be increasing in abundance, while sugar maples are in decline (Fei and Steiner, 2007; Oswald et al., 2018). But how can these patterns be generalized across the remainder of the genus? Will those species that already live in warmer and drier climates be favored by our warming and increasingly drought-prone anthropogenic climate? Or are species from cooler, wetter habitats secretly concealing a capacity to put up with a wider range of conditions than indicated by their current distribution? In my ongoing study of the climate-change vulnerability of the maples, I seek to answer these questions. Most temperate forests will experience hotter conditions and a greater risk of drought as our climate changes. What will this mean for the maples? In comparisons of diverse woody plants from across the globe, a few physiological traits have emerged as excellent predictors of how good a given species is at thriving in hot, dry conditions. One of these is turgor loss point, the water potential at which a leaf from a tree or shrub loses turgor, or wilts (Bartlett et al., 2012). To date, I have measured turgor loss point for seventeen species of maple coming from diverse sections of the genus and from all over the world. One pattern emerging from these data is that European and West Asian maples have the lowest (most drought tolerant) turgor loss points, followed by North American and then East Asian species. It appears that species living in the genus's original homeland are among the most intolerant maples of hot, dry conditions. This could be due to differences in the climatic Model Maples 13 histories of the maples' East Asian, North American, and European ranges. In Europe, for instance, cycles of glaciation and warming have likely pushed to extinction any species that could not survive dry conditions; East Asia likely contained more refugia, allowing these species, including many maples, to survive to the present (Tiffney and Manchester, 2001). Yet the perils of hot and dry conditions are not the only challenge that climate change will pose to the temperate maples. Paradoxically, some temperate trees may be at greater risk of springtime freezing damage in a warming climate. The increasing likelihood of false spring events, in which warm, short winters allow plants to begin growing again earlier in the year, may lead some trees to lose cold hardiness and sustain critical damage from sudden drops in late winter temperatures. To understand whether maples will be sensitive to false springs, I partnered with my colleague and fellow postdoctoral Putnam Fellow Al Kovaleski, an expert in measuring cold hardiness in woody plants. In these tests, we generally find that maples can withstand much colder temperatures than they are likely to experience in their native habitats\u2014or in Boston! Many of our test species, even when actively growing, flowering, or putting out new leaves for the spring, can withstand freezing to 14\u00b0F (-10\u00b0C) or below. It seems unlikely that false springs concomitant with a warming climate will expose trees to these temperatures at the right time of the year and for the periods of time necessary to cause considerable damage. However, in our analysis of cold hardiness across species, we found a continental pattern that echoed my work on tolerance of drought. East Asian species were, once again, least tolerant of cold conditions. But, in contrast to intercontinental differences in turgor loss point, North American species were generally the most cold hardy, with European species intermediate and East Asian species most vulnerable. This is likely due to differences in the way that each continent experiences the onset of spring. Recent In the author's research on drought tolerance and cold hardiness, Asian maples, like Acer ceriferum (right), tended to be the most vulnerable to severe conditions. European species, like A. campestre, tended to be more adaptable, especially to drought conditions. PHOTOS BY SUZANNE MROZAK 14 Arnoldia 78\/1 \u2022 August 2020 work by a group of ecologists led by Constantin Zohner (2020) has established that North American forests have historically been much more likely to experience freezing temperatures in late spring. European and Asian floras have responded to generally more permissive springs by developing an opportunistic strategy, taking more risks and initiating growth earlier in the spring. As a result, European trees are thought to be in the most danger of damage from false springs as the climate continues to warm. Back to the Future But what do these findings mean for the climate-change resilience of maples and, by extension, other temperate woody species? For me, the best way of thinking about the future is to once more turn to the genus's past. For instance, though it's impossible to measure the physiological diversity of the historic European and North American maple floras, we might imagine that, under warmer and wetter conditions, in which environmental pressures were laxer, each region was home to maples adapted to a variety of environments. But paleontological evidence, climate modeling, and the present-day existence of only drought-tolerant European maples and relatively cold-tolerant American species suggest a compelling story. Successive cycles of cold, dry, and glaciated ice ages during the Oligocene, interspersed with warming, may have weeded out those species sensitive to these environmental stressors. This mechanism has been suggested to explain the relative lack of diversity in temperate eastern North American and Eurasian forests relative to those in East Asia and western North America (Qian and Ricklefs, 2000; Svenning, 2003). And they certainly explain patterns of physiological and species diversity for the extant maples outside of East Asia. Perhaps the few remaining species in North America and Europe are those that have managed to survive, adapt, and migrate in response to glaciations and accompanying cold and dry conditions. Pushed into Meso- and Central America and into Northern Africa, they have subsequently returned to higher latitudes, becoming locally abundant in the case of widespread species such as sugar, red, Norway, and sycamore maples, among others. Differences in the geography and climate of each continent have only reinforced these adaptive patterns: North America, for instance, has more extreme and variable early spring temperatures, producing a more coldhardy flora (Zohner et al., 2020). But what will happen next, as our climate warms and patterns of precipitation become more erratic? We could be facing a future reminiscent of our past, but with a twist. Paleoecologist Kevin Burke and colleagues, in 2018, offer a particularly compelling illustration of how global change might affect Earth's temperate forests. In this modeling exercise, the authors compare likely scenarios of future climate change to what we know about the Earth's historical climate based on climatological reconstructions. Disturbingly, they predict that, under a business-as-usual scenario, in which humans do nothing to curb climate change, the Earth very well may, by 2200, have a climate akin to that of the Eocene period (roughly fifty million years ago in their analysis). This means that, in just seven maple generations, we may skip over fifty million years of changing climate, reverting to conditions similar to those before the diversification of the maple clade in the late Eocene and early Oligocene. Whether our current palette of maple species can tolerate these conditions is unclear. Our warming climate could open up some habitat, at least initially, for cold-tolerant and northerly distributed maples (like sugar maple and Norway maple) to extend further into areas that are presently too cold for trees to inhabit. And maybe the highly drought-tolerant species of Eurasia will be able to capitalize on drying and warming conditions to displace moremesic species. But there is also a likelihood that anthropogenic climate change will create what climatologists call no-analog conditions, a climate unlike one that our existing flora and fauna have ever experienced, much less adapted to. In this case, humans will truly face something new under the sun, as will our biotic companions. In the immediate future, the story of the maples suggests the critical need to begin the conservation of woody plants we know to Model Maples 15 be heat- or drought-intolerant. These conservation efforts include the ex situ migration of species out of their current native ranges. We also should begin investing in forestry that focuses on those species that, by virtue of their natural history, have proven themselves capable of withstanding long periods with limited access to water. These suggestions, based on the climatechange vulnerability of the maples, are meant to apply to a variety of temperate woody taxa, including oaks, willows, and birches. Each of these genera is the result of a complex set of journeys across multiple continents and survival over many millions of years of global change. That such adaptation is possible should at least give us hope for the work ahead of us to keep the planet livable, both for our own progeny and for whatever comes next in the story of the Northern Hemisphere's forests. References Bartlett, M. K., Scoffoni, C., and Sack, L. 2012. The determinants of leaf turgor loss point and prediction of drought tolerance of species and biomes: A global meta-analysis. Ecology Letters, 15: 393-405. Coiro, M., Doyle, J. A., and Hilton, J. 2019. How deep is the conflict between molecular and fossil evidence on the age of angiosperms? New Phytologist, 223: 83-99. Fei, S., and Steiner, K. C. 2007. Evidence for increasing red maple abundance in the eastern United States. Forest Science, 53: 473-477. Gibbs, D., and Yousheng, C. 2009. The red list of maples. Richmond: Botanic Gardens Conservation International. de Jong, P. C. 1994. Taxonomy and reproductive biology of maples. In D. M. van Gelderen, P. C. de Jong, and H. J. Oterdoom (Eds.), Maples of the world (pp. 69-99). Portland, OR: Timber Press. Li, J., Stukel, M., Bussies, P., Skinner, K., Lemmon, A. R., Lemmon, E. M., Brown, K., Bekmetjev, A., and Swenson, N. G. 2019. Maple phylogeny and biogeography inferred from phylogenomic data. Journal of Systematics and Evolution, 57: 594-606. Oswald, E. M., Pontius, J., Rayback, S. A., Schaberg, P. G., Wilmot, S. H., and Dupigny-Giroux, L. A. 2018. The complex relationship between climate and sugar maple health: Climate change implications in Vermont for a key northern hardwood species. Forest Ecology and Management, 422: 303-312. Qian, H., and Ricklefs, R. E. 2000. Large-scale processes and the Asian bias in species diversity of temperate plants. Nature, 407: 180-182. Svenning, J.-C. 2003. Deterministic Plio-Pleistocene extinctions in the European cool-temperate tree flora. Ecology Letters, 6: 646-653. Tanai, T. 1983. Revisions of Tertiary Acer from East Asia. Journal of the Faculty of Sciences, Hokkaido University, 20: 291-390. Tiffney, B. H., and Manchester, S. R. 2001. The use of geological and paleontological evidence in evaluating plant phylogeographic hypotheses in the northern hemisphere tertiary. International Journal of Plant Sciences, 162: S3-S17. Walther, H., and Zastawniak, E. 2005. Sapindaceae (Aceroideae) from the late Miocene flora of Sos'nica near Wroc\u0142aw - A revision of G\u00f6ppert's original materials and a study of more recent collections. Acta Palaeobotanica, 45: 85-106. Wolfe, J. A., and Tanai, T. 1987. Systematics, phylogeny, and distribution of Acer (maples) in the Cenozoic of Western North America. Journal of the Faculty of Sciences, Hokkaido University, 22: 1-246. Zohner, C. M., Mo, L., Renner, S. S., Svenning, J.-C., Vitasse, Y., Benito, B. M., Ordonez, A., Baumgarten, F., Bastin, J. F., Sebald, V., Reich, P. B., Liang, J., Nabuurs, G. J., De-Migueln, S., Alberti, G., Ant\u00f3n-Fern\u00e1ndez, C., Balazy, R., Br\u00e4ndli, U. B., Chen, H. Y. H., Chisholm, C., Cienciala, E., Dayanandan, S., Fayle, T. M., Frizzera, L., Gianelle, D., Jagodzinski, A. M., Jaroszewicz, B., Jucker, T., Kepfer-Rojas, S., Khan, M. L., Kim, H. S., Korjus, H., Johannsen, V. K., Laarmann, D., Langn, M., Zawila-Niedzwiecki, T., Niklaus, P. A., Paquette, A., Pretzsch, H., Saikia, P., Schall, P., Seben, V., Svoboda, M., Tikhonova, E., Viana, H., Zhang, C., Zhao, X., and Crowther, T. W. 2020. Late-spring frost risk between 1959 and 2017 decreased in North America but increased in Europe and Asia. Proceedings of the National Academy of Sciences of the United States of America, 117: 1-9. Jake J. Grossman is a visiting assistant professor of ecology at Swarthmore College and a research associate at the Arnold Arboretum. From 2018 to 2020, he was a Putnam Fellow at the Arboretum."},{"has_event_date":0,"type":"arnoldia","title":"Essential Gardening: Public Gardens in the Spring of COVID-19","article_sequence":3,"start_page":16,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25710","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24ea36c.jpg","volume":78,"issue_number":1,"year":2020,"series":null,"season":null,"authors":"Forrest, Todd; Galligan, Brian; Gott, Ryan C.; Guidarelli, Conor; Henrichsen, Esther Truitt; Huang, Terry; Kartes, Nancy; LaPlume, Greg; Loving, Sharon; Merriam, Debbie; Salyards, Jim; Shearer, Kim; Williams, Kevin","article_content":"The spring of 2020 has been defined by upturned plans. As the number of COVID-19 cases skyrocketed, lives across the United States were reconfigured. Eight-year-olds learned how to take school classes remotely. Grocery stores began limiting the number of shoppers who could be inside at once. Restaurants reinvented their menus for takeout. Businesses large and small closed their doors, sending millions of workers to unemployment. Even hospitals\u2014while stretched beyond the max on one front\u2014began furloughing employees, given that routine and elective appointments were canceled. Streets in cities like Boston became veritably empty, with no morning rush, no evening rush. Public gardens, like other cultural institutions, were confronted with the same stay-at-home mandates that shuttered their communities. According to the American Public Gardens Association, more than 25 percent of gardens closed on a single day (Monday, March 16), and by the end of March, only 4 percent remained fully open to the public. The plants, of course, did not wait to begin growing until gardens reopened. The sunshine-colored blossoms of forsythia and daffodils put on their radiant shows no matter what. The unrelenting arrival of spring was, in many ways, incongruous with the national mood. It also meant that horticulturists at public gardens continued working despite closures and event cancellations at their institutions. Schedules changed. Procedures changed. But there were plants to be tended. Below, thirteen horticulturists from gardens around the country describe the on-the-ground realities of car- Essential Gardening: Public Gardens in the Spring of COVID-19 Todd Forrest, Brian Galligan, Ryan C. Gott, Conor Guidarelli, Esther Truitt Henrichsen, Terry Huang, Nancy Kartes, Greg LaPlume, Sharon Loving, Debbie Merriam, Jim Salyards, Kim Shearer, and Kevin Williams ing for their collections during the first months of the pandemic\u2014the months in which an old normal faded and a new normal was created. January 24 On January 21, 2020, our nation's first case of COVID-19 was reported in the Seattle area, just a few miles from Bellevue Botanical Garden. I was not paying attention to the news. As garden manager, I was deep into planning our first big event of 2020: a Lunar New Year Celebration scheduled for February 2. We had been snowed out the year before, which would have been our first year celebrating this event, so excitement was high over the favorable weather forecast. We expected over one thousand visitors. I could imagine red-and-gold lion dancers snaking through entry gardens that would be redolent with witch-hazels and sarcococca. The hot pink blossoms of Camellia 'Mary Christian'\u2014evocative of the tea plant (C. sinensis)\u2014would be punching through the winter gray. One of our community partners, Lily, began each planning meeting by serving different varieties of Chinese tea. Her gracious habit kept me connected to the mission of our collaboration: teaching the public about botanicals used in Chinese teas. Lunar New Year was to be the first of four events celebrating Chinese tea arts through the seasons. At our pre-event check-in on January 24, Lily was visibly shaken. She was wracked with concern over the news out of China. She had been in touch with friends and family there and felt it would be disrespectful to hold a large public celebration at a time when so many were suffering. She and her colleagues feared the virus would spread here. I Facing page: Horticulturist Jessica Kaplan cares for the New York Botanical Garden's Native Plant Garden. PHOTO BY THE NEW YORK BOTANICAL GARDEN FORREST, T., GALLIGAN, B., GOTT, R. C., GUIDARELLI, C., HENRICHSEN, E. T., HUANG, T., KARTES, N., LAPLUME, G., LOVING, S., MERRIAM, D., SALYARDS, J., SHEARER, K., AND WILLIAMS, K. 2020. ESSENTIAL GARDENING: PUBLIC GARDENS IN THE SPRING OF COVID-19. ARNOLDIA, 78(1): 16-31 18 Arnoldia 78\/1 \u2022 August 2020 agreed to cancel the event out of respect for the Chinese community. At the time, I didn't think it was necessary to add \"out of an abundance of caution.\" It was a scramble to put the brakes on with just a little over one week's notice: cancel the lion dancers, the musicians, the tea ceremonies. Notify the public, the volunteers, the dignitaries. While our garden was deep in winter dormancy, with so much unseen beneath the surface, novel coronavirus was silently making its way through our community. Our area was destined to be the first epicenter of the outbreak in the United States. Events and programs fell like dominos, one after the other as our understanding of the pandemic evolved, until our governor issued a stay-at-home order and everything ground to a halt on March 25. Our facilities closed. A handful of crew members would continue coming in to care for the garden, which remained open for walking, free as always. Everyone began panic shopping for toilet paper, which I could not understand. I stocked up on veggie seeds and compost. On January 24, I didn't see any of that coming. I now feel haunted by that day, by my ignorance in thinking that the virus was far away, not our problem. Thanks to our Chinese friends, we made the right call and that decision may have saved lives. I remember that we, at Bellevue, are the lucky ones: no staff layoffs, volunteers eager to return, and all of them healthy. I remember that we are strong and resilient. And I remember that, in the garden, the hidden promise of winter dormancy burst into an early spring, with daffodils, daphne, and rhododendrons coming into bloom, each, in their turn, providing respite from pandemic fears. \u2014Nancy Kartes, Garden Manager March 15 I was at home on Sunday, March 15, when we decided to close the New York Botanical Garden indefinitely to the public and nonessential staff to help prevent the spread of COVID-19. For several weeks prior to this decision, we had been following the news of the virus, communicating regularly with various government agencies, and planning for at least a partial shutdown. In spite of our preparation, the decision to close the garden at the height of our annual Orchid Show and on the eve of spring felt nonetheless sudden and severe. Even though we had dealt with temporary closures after 9\/11 and during various hurricanes and blizzards, none of us had ever experienced a long-term shutdown with no clear path to reopening. Gardens shouldn't close in spring. Starting at about seven o'clock that evening, I set out to call every member of the garden's horticulture team (nearly seventy people in all) to relay the news and to assemble a small crew to come in the next day. With 250 acres of designed gardens and curated plant collections and two glasshouses, the New York Botanical Garden needs tending every day. Fortunately, New York State deemed us an essential business, which made our horticulturists essential workers. Unfortunately, due to social distancing protocol and budgetary concerns, we could only bring in a reduced staff on an intermittent schedule. By ten o'clock, I had reached everyone and had confirmed twelve staff for the next day. Even as I grimly delivered the news of the closure to one colleague after another, I was buoyed by their hope and desire to help however they could. Hope in the face of COVID-19 was no small thing. At the time, none of us knew how severe the pandemic would become, but all of us were aware that New York City, with its densely packed humanity, could be fertile ground for a highly communicable virus. Many of my colleagues live in the five boroughs and take public transportation to the garden. Many are in close contact with elderly parents, or have partners with respiratory conditions, or care for small children, or have some other legitimate reason to be especially scared of contracting COVID-19. Despite their personal concerns, the horticulture staff knew what was at stake and gamely signed up to come in. They understood the essential nature of their work keeping the New York Botanical Garden healthy and beautiful so the garden can achieve its mission of serving and delighting the public. This spring has taught me a lot about the profound impact our garden has on peoples' lives. It has also deepened my respect and gratitude for the professional horticulturists who Public Gardens and COVID-19 19 care for our collections, displays, and natural landscapes. These are New York horticulturists: a bit jaded perhaps, and suspicious of authority (e.g., me), but as tough and serious as they come. Many have worked here for decades and are deeply proud of what we have built together on the foundation laid by the generations of horticulturists who came before us. And so, through the height of the pandemic, our now officially essential horticulturists came to the garden, albeit on significantly reduced and altered schedules, to care for our plants, COVID-19 be damned. \u2014Todd Forrest, Arthur Ross Vice President for Horticulture and Living Collections March 17 On Tuesday, March 17, I was in the nursery at the Arnold Arboretum with five other horticulture staff, digging trees and shrubs for the spring planting. It was the day after buildings at the Arnold were closed to non-essential personnel. Originally the closure was described as a \"trial\" work-from-home week, but for most, it would become a new normal. Horticulture staff would also set up home offices to reduce essential staff densities. Those who were juggling work, parenting, and teaching duties were home first. Our team in the nursery was the only horticulture and greenhouse staff on-site that day. The nurseries consist of three plots that are tightly spaced around the greenhouses, located on a central edge of the landscape. Visitors can see the nurseries and greenhouses through a chain-link fence. Almost all the one-hundredplus trees and shrubs that were slated to be dug this spring were growing in a single nursery plot, and everything was to be balled-andburlapped. This method involves hand-digging a teacup-shaped mass of roots and soil (the children's song \"I'm a Little Teapot\" always runs through my head) and lacing sisal in an inter- At the Arnold Arboretum, in-person horticulture meetings have shifted to Zoom. Here, horticulturist Conor Guidarelli attends from the landscape. JONATHAN DAMERY Public Gardens and COVID-19 21 woven drum-like fashion to hold on a covering of burlap. One person can do the digging and burlap dressing, but it takes at least two to hoist the plant to the surface. We managed to keep several feet away from one another by lifting with ball-and-burlap straps. At that point, we were all wearing makeshift masks fashioned from cotton rags and handkerchiefs. We normally dig several specimens and then go out onto the grounds to plant them on the same day. This spring, however, everything was to be dug at once. Given the small size of the nurseries, space is constantly in demand. New seed arrives from plant-collecting expeditions every year, and the seedlings work their way into the greenhouses, to the shade nursery, and eventually either into containers or into one of the nursery plots. If plants were not dug from the nurseries it would stop the production line. Because many of the plants are collected in the wild, throwing out the inventory is not a choice. The plants are impossible\u2014or exceptionally hard\u2014to replace. These are not massproduced Knock Out\u00ae roses. When the coronavirus was first being reported in the United States, back in January, I was grimly aware that the pathogen would be exceptionally disruptive. In our horticulture meetings, we began creating a game plan for how we would prioritize our operations under a series of scenarios. The fourth scenario was a near shutdown of operations, with only one or two people on-site. As it ended up\u2014and as I worried would happen\u2014we went straight from the modest precautions of the first scenario to the intense shutdown of the fourth within a matter of weeks. Before scenario four could occur, however, spring planting had to be completed. We had initially planned to tuck the balled-and-burlapped plants into another space in the nursery for a fall planting, but halfway through, the plan would shift: everything would be planted, including another two-hundred-odd plants that had been grown in containers. From the nursery, we could see through the fence to the main roadway that winds through the Arnold, which offered a view of a tremendous influx of visitors. The landscape is free and would remain open despite other closures. It is tucked directly within Boston residential neighborhoods, and with businesses around the city sending employees home\u2014or worse, laying off workers\u2014wave after wave of visitors were taking midday walks in the landscape. It felt like everyone was arriving for our largest event of the year, Lilac Sunday, but the lilacs were still more than a month from blooming. Tree branches were still bare and leafless. But our relentless pace of digging continued. This task was essential. \u2014Greg LaPlume, Arboretum Horticulturist March 25 In the early days of the pandemic, the pervasive singing of birds at Filoli was uncanny. The garden is nestled in the mountains between the San Francisco Bay and the Pacific, halfway between San Francisco and San Jose. While the gardens are formal\u2014part of a sprawling estate that was established on gold-mining profits more than a century ago\u2014wildlife is always abundant. Birdsongs provide a sense of vibrancy during the day, and large animals (like cougars, coyotes, foxes, and raccoons) leave evidence of nighttime visits. On March 25, I was in the Sunken Garden, snapping a social media photo of yellow 'West Point' tulips that were blooming within the low, clipped hedges of the parterres. The calls of sparrows, towhees, crows, and finches were inescapable, but they were now an eerie reminder of the lack of human voices in the garden. Filoli had closed the week before, on March 17, and although the horticulture team would continue to care for the landscape, the garden had to lay off some of our frontline staff at the beginning of our closure. Wildlife was becoming more brazen in their activities, but it was very bittersweet when all who would normally be enjoying the garden, along with the birds, were missing. Filoli has blooms 365 days a year because of the moderate climate along the coast of northern California. Camellias and daphne begin blooming in January. In summer, the formal parterres showcase a bounty of colorful designs. But spring continues to be our biggest draw. Locals and visitors from around the globe are captivated by the spring experience of seeing 'West Point' tulips at Filoli Historic House and Garden. PHOTO BY JIM SALYARDS daffodils and tulips in our meadows and formal beds. Wisteria clambers on the side of the mansion, and peonies are showstopping. But this year, our spring peak of mid-March to mid- April was completely missed. All the planting and tending on the part of the staff, all the expectant calls and emails that started at the beginning of the year asking the best time to visit were for naught. I did my best to share photos and videos through our social media outlets, but it's just not the same. A few thumbs-up or heart emojis are a poor substitution for the \"oohs\" and \"aahs\" and the thank-yous we receive from guests each day\u2014the guests who call out compliments while we are weeding and pruning or who pass along the praise to our colleagues in visitor services and interpretation. Public gardens like Filoli are champions of environmental education and conservation, yes, but we also provide substance for people's souls. Hopefully, in the near future, the garden will once again become a space of healing, just when the world needs us most. \u2014Jim Salyards, Director of Horticulture April 6 The first week of April, three weeks after Utah went on voluntary shutdown, I spent two beautiful days pruning at Ashton Gardens where I work. The gardens were closed to the public, and the gardeners were \"social distancing,\" a term that was new to our everyday vocabulary. I was on my own, pruning and listening to music within the walled Secret Garden, an enclosed space among the designed ruins of gothic arches that drip with honeysuckle (Lonicera japonica 'Halliana') and Virginia creeper (Parthenocissus tricuspidata). As the garden designer, I don't often have a chance to work directly in the At Ashton Gardens, a horticulturist cuts tulips from the Italian Gardens for online sales. ESTHER TRUITT HENRICHSEN Public Gardens and COVID-19 23 gardens, but during the pandemic, the entire staff was stepping up to help with horticultural care. I love to prune. I worked my way through the collection, shaping and thinning shrubs. I pruned branches from an indecisive willowleafed pear (Pyrus salicifolia 'Pendula'), removing those that were sticking straight up and keeping those that were draping down. It was just me, the blue skies and sunshine, and ten thousand tulips in the Secret Garden\u2014yes, ten thousand of the quarter million that we had planted at Ashton Gardens for our annual Tulip Festival. Like so many things this spring, the event did not happen, so only the gardeners and the birds were there to see the show. Ashton Gardens lies thirty miles south of Salt Lake City in the foothills of the Wasatch Range. We are part of Thanksgiving Point Institute, a complex of gardens, a farm, and museums in the middle of a rapidly developing area called Silicon Slopes. The Secret Garden is one of our guests' favorite spaces within our fiftyfive- acre landscape, and its charm lends itself as a place to stage marriage proposals, as well as for meetings of the local crochet club. There were no marriage proposals in the gardens this spring. Utah went on voluntary lockdown on March 14, before the gardens officially opened for the season. The office staff of Thanksgiving Point mostly worked from home, and staff with service jobs were paid to stay home. Due to our ability to distance ourselves while working, the garden staff had the good fortune of continuing to work every day. On March 18, four days after lockdown, a 5.7-magnitude earthquake hit the Wasatch Front. I was in my office and, true to my elementary school earthquake-drill training, dove under my desk. The staff working outside in the gardens didn't even feel the quake. Over the next several weeks, over two thousand aftershocks occurred, and COVID-19 was always there just beyond the garden gates. While we worked in the gardens, nature helped us to find solace in the storm and feel less unnerved than many in our community. Throughout the weeks that would have been the Tulip Festival, we cut fifty dozen tulips from the garden each day. Thanksgiving Point members could preorder bouquets and have them delivered to their front porch. Or they could drive past the Ashton Garden Visitor's Center, pop the hatch so that garden volunteers could put the flowers into the car, and then drive away with their little bit of the Tulip Festival. \u2014Esther Truitt Henrichsen, Garden Designer April 13 On Monday, April 13, I was at the Mary May Binney Wakefield Arboretum, located just south of Boston. As the arboretum director but one of just four employees, I normally work in our gardens every day. It's a twenty-five-acre property, listed on the National Register of Historic Places, and we are recognized for our collection of more than three hundred kousa dogwoods (Cornus kousa). The dogwoods were grown from seed that Mary (Polly) Wakefield collected at the Arnold Arboretum, where she took propagation classes for more than forty years. Our grounds had officially closed to the public following the statewide stay-at-home order that went into place on March 24. Due to the size of the landscape and staff restrictions\u2014 one week on and one week off\u2014I had only seen my coworkers in passing and felt so fortunate to still be employed, working in the garden that had become such a place of peace and solitude. On that day, however, pouring rain kept me inside catching up on paperwork. In an effort to create distance between staff members, I had moved my office from an old farmhouse to the main historic residence: a 1794 Georgian mansion. As the rainy and blustery spring day progressed, I looked out of the windows of the antiquated kitchen where I had set up my workstation and observed several of our massive hemlocks swaying back and forth. Mature trees before the Civil War, these hemlocks withstood the 1938 hurricane that destroyed many native trees all over New England. More recently, these giants survived hemlock woolly adelgid with attentive care. As winds strengthened, I received an alert from the Blue Hill Observatory, just a mile away, that gusts were recorded at eighty miles an hour. I heard a loud crack and saw that a large limb had come down and obliterated our ten-foot privacy fence. I stepped out of the mansion to survey the damage and quickly realized it was not a good time to be 24 Arnoldia 78\/1 \u2022 August 2020 standing among so many towering trees: hemlocks, sugar maples, and red oaks that are the oldest in our collection. Instead, I locked up the mansion, hopped in my car, and headed toward the gate and home. Ten minutes later, the largest hemlock snapped about ten feet up and with its huge girth took another massive tree with it, narrowly missing the mansion. I received an alarmed text from my colleague saying, \"We lost the big ones.\" I could not fathom these trees falling until the images appeared on my phone. It seemed almost cruel that these hemlocks would no longer record history from their stately position. As a wave of sadness came over me, I recalled a moment standing in an ancient hemlock grove in graduate school, listening to Mark Ashton, my favorite forestry professor. He spoke with deep passion and amazement about the hemlock's ability to survive in the understory for hundreds of years, biding their time until one of their cohorts comes down leaving growing space for a young tree to continue the legacy. Perhaps this would be an opportunity to plant some of the Chinese hemlocks (Tsuga chinensis) that I had been raising in our nursery. Or perhaps a young self-sown hemlock seedling would rise within the gap and thrive. I took great comfort in the fact that plants are so resilient and will go on, as will we in this challenging time. \u2014Debbie Merriam, Arboretum Director April 15 One morning, around April 15 (time has lost a lot of meaning during quarantine), I came into work at Phipps Conservatory and Botanical Gardens to a desk covered with plant samples. As the associate director of integrated pest management, I handle our plant health care, including the diagnosis of all pests and disorders and the prescription of management and scouting protocols. I typically expect interesting new mysteries to appear. But looking through these samples, my first thought was, \"These are all known issues. Why were they turned in?\" Then, it hit me. Of course. With our new coronavirus-altered schedules, horticulturists were caring for spaces and plants they never had before. Everything they encountered was new to them\u2014the day-to-day simply didn't exist anymore. While gardens may not have been deemed officially life-sustaining during quarantine, we certainly are in the business of sustaining life\u2014 plant life that is. Phipps, located in Pittsburgh, closed to the public on the afternoon of March 14 due to COVID-19. While our glasshouses\u2014 a mixture of original Lord & Burnham Victorian- style rooms from 1893 and some modern additions like our Tropical Forest Conservatory\u2014 were shuttered to visitors, our plants still needed attention. Before coronavirus, horticulturists managed specialized areas including production greenhouses and collections of palms, ferns, orchids, cacti, bonsai, and more. During the coronavirus closure, we had large changes to our team structure. Instead of furloughing staff, we reorganized our horticulture department into three small teams that rotated on-site coverage on continuous five-day schedules starting March 27. All other Phipps staff worked from home, including horticulturists during the off-site portion of their rotations. This meant horticulture staff cared for more spaces than they typically would, often outside of their plant specialty. Pieces of our pest management plans, such as syringing this or that palm to suppress spider mite populations, were not always tended to since staff were working hard to perform basic plant life support like watering over large areas. These sorts of pest management tasks that were second nature to a room's usual horticulturist were unknown to other staff. In normal times, a handful of volunteers are at Phipps nearly every day, helping horticulture staff pot plants, clean beds, sow seeds, and more. One volunteer comes in every week to help me by scouting greenhouses for pests, carefully washing and cleaning plants to manage insect issues, and collecting samples of leaf spots and rusts. Like the other volunteers, she has not been on-site since we closed to the public, and without her, these simple but time-consuming tasks fell by the wayside. On May 3, according to my notes, I was finally able to check a particular cycad that my volunteer would regularly clean but that hadn't received her attention in almost two months. Public Gardens and COVID-19 25 The poor plant was overwhelmed with mealybugs, its newest set of growth fuzzy with wax and deformed and stunted from the mealybugs' toxic saliva. I cleaned it, arranged for augmentative biocontrol releases, and even identified some wonderful brown lacewing predators that were already present, feasting on the mealybugs. I, and all our staff and plants, are really missing our volunteers. Integrated pest management and plant health care more broadly are team efforts. As we adapt to whatever our new daily \"normal\" will look like, I've come to see how cross-training staff in many areas would produce knowledge and skill redundancy in all aspects of plant health care. Colleagues at other gardens have had similar realizations. These and other conversations and innovations will move gardens forward and better prepare us\u2014and our plants\u2014for whatever the future may hold. \u2014Ryan C. Gott, Associate Director of Integrated Pest Management April 20 The headhouse sits at the southern end of Filoli, among the greenhouses, nursery, and a few oaks. Inside, the air was cool and faint with the soft scent of aged cement and redwood. Working in the dim light, I slowly organized my desk. My fellowship had ended early due to the pandemic, and this was my final day. I sorted through the years of accumulation drawer by drawer, encountering fragments of the many lives that had passed through here: a handwritten reminder, a hair tie, a playing card, a dead spider. My mind drifted as I worked. I had been working off-site for a month and a half, and my last memory of the garden was in early spring. The hellebores and magnolias had just given way to a few blousy spring camellias, but most of the garden still slumbered. While my life took a pause, the strengthening sun and late spring rains had coaxed the garden out of its winter dormancy. Now, the fresh green growth of redwoods, coastal oaks, and arbutus enrobed the Santa Cruz Mountains. Irises and tree peonies stretched their satiny crepe petals in the spring sunlight. Masses of tulips swayed cheerfully in the gentle breeze as voles darted between their beds. With so many flowers in bloom and no one to admire them, the garden was rejoicing, blooming for itself without judgement. A little space to breathe, a moment to grow. After labeling the stacks of important documents and wiping down all surfaces, I headed out to the staff vegetable garden. Tucked away behind the headhouse, the garden is protected by a tall cherry laurel hedge and brick wall. The winter crops had finished. I saw evidence of recent activity, but not a single soul was there. Future plots were weeded, tilled, and enriched. Rows were marked and irrigation laid waiting. Soon rows of tomatoes would glisten in the sun, their leaves releasing a resinous fragrance. Swollen squash would hide under their giant prickly leaves. Multicolored carrots and potatoes would be unearthed like crystals and geodes, while sun-warmed strawberries and bright lemon verbena perfumed the air. The abundance would provide more than enough for human, beast, and microbe. For me, this was a place of refuge that had sustained me for a year, a place where I cultivated community with the earth and between people. I will miss the way the soil crumbled in my hands and how laughter floated over the garden hedge. Walking down the gravel path one last time, I took in the peace before heading out through the garden gate. \u2014Terry Huang, now Assistant Director of Mildred E. Mathias Botanical Garden at UCLA April 30 I dragged my eyes away from yet another Excel spreadsheet to the tall casement windows in my office. From my vantage at the northwest corner of the Horticulture Building at Longwood Gardens, I could see threatening clouds lumbering towards me. Located forty miles southwest of Philadelphia, Longwood doesn't normally receive such severe weather in late April. The sky darkened in minutes and began hurling something between hail and enormous raindrops into the windows, blurring my view of the forest edge across the deserted employee parking lot. The forecast was for a straight-line wind, but the tree branches were swirling in circles. The tops of the tall oaks and tulip poplars swayed in an unnatural dance. It was late on April 30, and six weeks had already passed since the mandatory closure 26 Arnoldia 78\/1 \u2022 August 2020 of our doors. Like the closures at many other gardens, this prolonged shutdown has been a first for Longwood. Even after a long career, I had found myself making the most challenging decisions I have ever had to make\u2014reducing budgets, furloughing all of the part-time staff, preserving our precious and rare plant collections\u2014 all within the span of a few weeks. We had already removed thousands of plants slated for our spring display, ripping out hundreds of beautifully planted beds of tulips and lilies. Our greenhouse staff had meticulously grown Echiums for a full year, and although the plants were magnificent, towering at four feet high, they never made it to display but instead were diverted to our compost stream, along with thousands of other crops. For a gardener, shuttering such beauty is heartbreaking. I hadn't expected to have such an emotional response to all of this (after all, they're only plants). Musing about the challenges being faced by our communities and our nation, I chided myself for being selfish. Our losses were only short-term. Watching the wind and rain, I suddenly heard a loud, splintering crack followed by an earthshaking crash. I squinted through the gray deluge and could see an enormous oak was missing from the skyline. Shortly after, the phone rang and a colleague delivered more bad news: a precious tree, the oldest Magnolia acuminata var. subcordata 'Peirce's Park' in the garden, named for the historic site on which it was planted, was down. This cucumber magnolia, at over eighty-fivefeet tall and with a mighty girth of four-feet, was a national and state champion. We had traced the lineage directly back to French explorer Andr\u00e9 Michaux, who discovered this species in South Carolina in 1788. Planted between 1780 and 1830, this tree was part of a mature alle\u00e9 of trees (considered to be the \"Soul of Longwood\") that the Peirce brothers had established on the property. Garden staff have even speculated that John Bartram or William Hamilton had helped them procure the tree, as they had in their own gardens in Philadelphia. Our founder, Pierre S. du Pont, purchased the original farm that was to become Longwood to save these trees from being lumbered. It was too dangerous to check the tree that night. The following morning, breezy and clear skies laid bare the torn twigs and stripped leaves strewn across the paths and lawns. I kept walking; I have become better at ignoring the lack of perfection these past months. Normally the entire horticulture team would be scouring the landscape picking up every bit of debris\u2014but not today. Seeing the tree down reminded me of an image of a poached elephant I had seen years ago: gray, lifeless, enormous, and sprawled awkwardly across the path. I ran my hands over the tree's scaly bark, gave it a hug, and said goodbye. On my way back, grief gave way to a fleeting thought of hope. Almost twenty years ago, we propagated saplings from this tree. I walked past three of its progeny that were battered and dazed but still standing strong: future sentinels at the opposite end of the same alle\u00e9 the Peirce brothers planted more than two hundred years before. \u2014Sharon Loving, Vice President of Horticulture May 8 On Friday, May 8, I received a text message from Erin Bird, our communications manager. \"Are the lilacs blooming?\" she asked. I knew she was looking for social media content. It was a welcome text. I missed these interactions. I hadn't seen or heard from many of my coworkers since the Denver Botanic Gardens closed on the afternoon of March 16. Thankfully, none of my colleagues lost their jobs, but only staff considered essential to the stewardship of the facilities and living collections were on campus, so the space felt different, quiet, empty. \"They are blooming, but not very well \u2026\" I responded. Denver sits in the rain shadow of the Rocky Mountains, on the eastern edge of the North American steppe, the expansive semiarid grasslands and shrublands characterized by hot summers, cold winters, mineral soils, and very little water. Echoing the mountains that reign above the plains, this extreme environment lends itself to extreme weather. Long stretches of warm weather start in February, encouraging early leaf growth and bud break, only to be followed by sharp freezes in late April. These fluctuations are particularly hard on introduced flora during spring and fall, when biorhythms can be out of sync with steppe weather. Due to one 14\u00b0F (-10\u00b0C) day in April, this year had turned out to be a particularly disappointing season for the lilac (Syringa) display. Still, there were some blooms. What had survived was fragrant and beautiful. And I realized what Erin must have already been thinking about\u2014 that for the first time in almost seventy years, we couldn't directly share this experience with our community. And although almost everything human was static, the phenological rhythm of the lilacs exists beyond COVID-19 time (and our psyches) as heralds of change and expectation. \"I'll go get some pics for you.\" I moved through the collection, across the pathways of fescue, yarrow, clover, and flowering daisy, photographing the most hopeful, vibrant panicles. It was quiet, and I was alone in the collection at a moment when these shrubs would normally be stroked, sniffed, and praised thousands of times a day. I felt selfish and wondered if the lilacs were suffering from a lack of attention. They are gregarious creatures, attention seekers, but still, they felt dull: their scent weakened by my mask and their presence hazy through my constantly fogged sunglasses. Sometimes I feel comforted by our new personal protective equipment, but at that moment, I felt smothered and separated. The standard protection that comes with being a gardener is, in itself, sometimes bulky, sometimes tactically empowering: hiking boots and knee pads for constant squatting, crawling, and walking; gloves to stop the earth from tearing my hands apart; and sunscreen, sunglasses, and a wide-brimmed hat, all to fight off the extra mile of solar radiation that we get in the high plains. The addition of the cloth mask was reassuring at first, during the colder days of March and early April, but on that hot May afternoon, it felt oppressive. Lilacs (Syringa vulgaris 'Henri Martin') at the Arnold Arboretum. JONATHAN DAMERY I finished photographing the blooms, put my phone in my pocket, and walked deep into a bed, hiding from the ever-present gaze of the mountains. I took my gloves off, pulled my mask down, and pushed my face in to fully smell their chemistry. To breathe together, to share the air with plants, has always been our most fundamental of exchanges. \u2014Kevin Williams, Horticulturist May 8 On the Friday before Mother's Day, I was working in the lilac collection at the Arnold Arboretum, double-checking it for any missed deadwood, overlooked weeds, and spillover mulch. As the caretaker for the collection, the upcoming Sunday would have been day zero for me\u2014Lilac Sunday\u2014the moment when all the work that I do comes to a head. Lilacs are the only plant that gets its own day of festivities at the Arnold, but unlike the 111 years of prior celebration, this year's event had been called off. The lilac collection is nestled on one side of Bussey Hill, which rises in the center of the landscape. In early May, the collection looks endless as it wraps its way along the contours of the slope, with bursts of violets, purples, pinks, and whites. Normally, on Lilac Sunday\u2014 a Mother's Day tradition\u2014thousands of New Englanders clamor to get their photo taken with the prolific flower displays and relish in the sweet aroma. Tour groups pack the roadway, enthusiastic and inquisitive. Merchandise and information booths are spaced accordingly. In recent years, an ensemble from the Boston Symphony Orchestra performed as guests made their way among the shrubs. But all the pomp and circumstance for this year was scrapped in light of the pandemic. I felt confused in my plight to steward the collection with all the fear and uncertainty that hung heavy in the community. Given that the Arnold Arboretum is one the few gardens to remain open while the rest of the world sheltered in place, I continued with \"business as usual,\" so that some semblance of \"normalcy\" might be evident to any visitors who still relied upon the lilacs for their spring awakening. The Friday before is usually a buzz of activity for me as I coordinate seasonal employees, interns from a local agricultural high school, and fellow colleagues to assist me in the final touch-up and presentation. I'm frequented with questions from the public about tips and tricks for lilac care, but this time I was isolated in my work. The hustle felt more imminent this time, not only because of the lack of extra hands but also because the preparations had taken a different slant. I always pamper and cater to the plants. I try my best to not distort a shrub's natural growth habit, but this time, my focus had changed to looking for pinch points in the collection. Where neighboring bushes might be funneling individuals too closely, I began pruning aggressively to widen corridors for greater social distancing, should the people we had asked to stay home decide to visit after all. I also spray-painted white arrows on the sidewalk to request one-way traffic to limit potential exposure of those in the garden. I spent the afternoon posting normal signage (\"Please don't pick the lilacs,\" \"No picnicking at the Arboretum\"), along with another, \"Don't smell the lilacs.\" It felt strange and unfounded, especially for someone accustomed to removing hazards or providing a safe environment for visitors. But treating the lilacs like they could transmit the coronavirus was the necessary precaution given all the uncertainty. The day was a complete fog for me, literally, as I ran around with clouded safety glasses from my mask. My instincts drew me in to pull the lastminute weeds and to cut out the hidden deadwood, but my main directive was to make visitors aware of the unseen dangers of what had always been a joyous day for celebrating spring, mothers, and the season of brighter days to come. \u2014Conor Guidarelli, Arboretum Horticulturist May 12 On the morning of May 12, I walked rows of Magnolia hybrids at the Morton Arboretum, investigating buds on the trees, searching for signs of life. My work as the tree and shrub breeder at the arboretum builds on the legacies of others. I am less than four years into my career, yet I have tree selections in the pipeline and populations of progeny to select from. 28 Arnoldia 78\/1 \u2022 August 2020 Public Gardens and COVID-19 29 These selections are the culmination of almost a hundred years of work spread over the careers of many individuals. Without this team that came before me, I would not have been here in this field of magnolias staring at the freezeburned blossoms and emerging leaves. These hybrids had been developed over years by the late Dennis Ledvina, a much loved and highly respected magnolia breeder from the Green Bay area. Late-season freezes provide an opportunity to select for Ledvina's target traits\u2014 improved cold hardiness and delayed bloom time. The longer a tree holds off on developing its flowers or pushing out new growth, the better chance it will have of coming through these freezes unscathed. I pressed buds between my thumb and forefinger, flagging trees whose buds gave way with a satisfying squish. While many gardeners lament when a late freeze occurs, a breeder reviews a weather forecast and then sits in anticipation for these moments, grateful for the gift Mother Nature has bestowed. The orange flagging tape was my signal to wield the mighty chainsaw and give the tree one final prune, a single cut at the base. Not all was lost; some would make it to see another winter. My dog, Maybelle, ran up and down the rows, delighting in the freedom of being penned into a fifteen-acre nursery. The nursery\u2014wound round with electric fencing intended to keep out larger wildlife\u2014is located on a southern edge of the arboretum, nestled between part of the taxonomic collection and a berm that buffers the sounds of an interstate that races through Chicago's western exurbs. Maybelle stopped short, leaned down to the ground and tentatively sniffed, inhaling the traces of other animals not yet seen. My gaze followed Maybelle's: a female coyote stood thirty feet away, her belly hanging low and rippling with the life held within. After the Morton shut its gates on April 2, signs of coyotes have become widely apparent to anyone who accesses the grounds for essential work. The arboretum includes seventeen hundred acres of cultivated and curated collections, managed forests, and a planted prairie and savanna, which is more than enough room for coyotes and other wildlife. With only the familiar faces of arboretum staff present, these canine compatriots more readily emerge from their secret daytime hollows to observe us from Coyote at the Morton Arboretum. BEN CASCARANO a distance. While not overly comfortable with us humans, the coyotes always exhibit some curiosity toward our existence. Perhaps this is a legacy of a recently retired staff member who spent fifty years living and working on the grounds with his family. August members of the staff have widely shared a story, with eyes twinkling, about how, years ago, this staff member once shared hot dogs with the first coyotes to establish their pack on these grounds. Some decades from now, generations of magnolias and coyotes between, perhaps the greatgreat- great-granddaughter of this coyote will meet me in these rows one quiet spring day. As my thoughts wandered, I made eye contact with the coyote. She lingered momentarily, and as she walked away, I returned to Ledvina's magnolias. \u2014Kim Shearer, Tree and Shrub Breeder and Manager of New Plant Development Program May 15 On an exceptionally muggy day in mid-May, I paused to take in my surroundings in the vast tropical collections at Naples Botanical Garden. I stood on the boardwalk over the Water Garden, a favorite spot that offers a panoramic view of the 170-acre property. Our Smith River of Grass, the garden's central spine and a replica of the Florida Everglades, stretched in front of me. The jungle-like Lea Asian Garden arose on one side, and the bright green performance lawn extended on the other. The garden had never looked so lush, I thought. The staff had laid fresh mulch, pruned, weeded, planted, taken out ailing trees, cleared areas for new displays. The colors this morning were extra vivid, a visual effect brought about by soot blown in from a wildfire burning in the Picayune Strand, well to our east. The waterlilies glowed in the light\u2014mostly pinks, with a few yellows and purples. Beyond the pond, the cassia and poinciana lived up to their common names\u2014 golden-chained and flamboyant. The smell of smoke made my stomach turn. Disasters loomed in my mind\u2014the pandemic we are living through and the hurricane we survived in Florida not all that long ago. Irma tore a path through Naples in September 2017, shredding shrubs and downing trees. A local reporter described our garden as resembling \"layers of tossed salad.\" Our visual paradise, which we had created from barren swampland less than ten years before, was ruined. The View from the Water Garden at Naples Botanical Garden. PHOTO BY NAPLES BOTANICAL GARDEN Public Gardens and COVID-19 31 winds had blown away years' worth of sweat and dirty fingernails. I was wrong about the ruin. Volunteers and incredibly dedicated staff rushed to save the place. We rebounded like kudzu in Alabama. I'd argue we came back even better than before. When we first opened in 2009, we were a brandnew garden mostly focused on giving snowbirds something pretty to look at. We now have an incredible botanical collection, thanks to our amazing \"plant nerds\" and their desires for rare and unusual specimens. But there is a difference in this new disaster. This time it's not the plants but the team that is battered and bruised and beaten. The plants look amazing, and if we could invite guests back in, their experience could not be better. But our gates closed in mid-March, not only to visitors but to our volunteers and even to our families. During the hurricane recovery, our staff and volunteers took afternoon breaks together to cool off and enjoy lunch, a daily ritual that lifted everyone's spirits. Who could lift us now? I thought about how my team looked during one of our weekly staff meetings. We had been able to keep everyone on, and I knew they were glad to be working, but I could see their exhaustion. They're the \"tossed salad\" this time, I thought. It's not just the dirty shirts and the unshaven faces, it's a lack of purpose that I worried about. Everyone was giving 100 percent, but for what? No one could see our beautiful oasis. No volunteer force was going to stride in to relieve the workload and share in the joy of creating something special. But I shook off those feelings. We're adapting, just like nature taught us to do after Irma, when our collections rebounded and shone with the beauty I noticed that morning. We decided to take on big projects\u2014like transplanting trees and dredging ponds\u2014to give the team a break from the endless weeding and pruning and offer them the satisfaction of accomplishing major tasks. The gardeners, who are so used to interacting with the public, found other ways to communicate. They took photos, shot videos, and shared stories about the collections, broadcasting their work to the world online instead of welcoming guests to our property. They answered questions over Facebook instead of in person. Nothing can replace the in-person experience of a garden, but our horticultural creativity meant that all could share in the joy of something special\u2014even during a crisis. \u2014Brian Galligan, Vice President of Horticulture Epilogue By the end of the spring, gardens and arboreta began to reopen. Bellevue Botanical Garden and the Arnold Arboretum were among the few whose grounds remained fully open throughout the early months of the pandemic. Ashton Gardens reopened on May 1, allowing visitors to catch the late-blooming tulips, and Filoli reopened on May 11. Attendees at both gardens were required to purchase timed-entry tickets. Filoli initially offered eight hundred tickets each day and later raised the number to fourteen hundred. Prepurchased tickets became the modus operandi for gardens\u2014a way of preventing attendance surges and of reducing interactions between visitors and staff at entrance bottlenecks. Denver Botanic Gardens reopened with a ticketed entry on May 22. The Morton Arboretum reopened to members on June 1 and to the general public on June 15. Phipps Conservatory and Botanical Gardens reopened on June 13, allowing a one-way path through the indoor conservatories. Longwood Gardens reopened on June 18, about three weeks before a massive corpse flower (Amorphophallus titanum) came into bloom. Due to state-mandated limits on guest capacity, the garden significantly expanded their evening hours so that more visitors could obtain tickets to experience the rare and short-lived bloom. Some visitors were relieved to find that the notoriously foul smell of the flowers was muffled by their masks. Naples Botanical Garden fully reopened on July 6. New York Botanical Garden partially reopened on July 21. By the end of July, the Wakefield Arboretum had opened for limited reservation-only tours and special programs. The trajectory of the pandemic is far from over. Yet the innovations that have allowed gardens to reopen in person\u2014and to connect with visitors online\u2014will have a lasting impact, no matter what lies ahead."},{"has_event_date":0,"type":"arnoldia","title":"One Green Earth","article_sequence":4,"start_page":32,"end_page":41,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25711","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24ea76f.jpg","volume":78,"issue_number":1,"year":2020,"series":null,"season":null,"authors":"Raven, Peter H.","article_content":"While I was lying in bed in the spring of 1944, recovering from measles at the age of seven, my mother entered my bedroom and handed me a bright orange book: Six Feet by Ruth Cooper Whitney. Once I had taken a look, I couldn't put the book down. It presented simple stories, illustrations, and poems about different kinds of insects. Its stories were so engaging that I couldn't wait to rush outside and see the insects for myself\u2014 even though I'd paid no attention to them previously. In our garden, built on the sandy flats of the Richmond District in San Francisco, I could find and rear cabbage butterflies; discover Jerusalem crickets, earwigs, tenebrionid beetles, bumblebees, and other fascinating creatures; and begin to catch a glimpse of how they all fit together. I soon began making a collection of butterflies but then switched to beetles. I accumulated a collection of several boxes of these mounted insects, which my parents would proudly display when guests visited. As the years passed, it turned out that this book, an otherwise ordinary gift from my mother, provided the first major step towards my career as a botanist and environmentalist\u2014 a career that would culminate with a forty-year tenure as director of the Missouri Botanical Garden. During that career, I would see our global understanding of biodiversity expand far beyond what was known when I first began collecting insects in my childhood backyard. Yet, over that same period, researchers have shown how humans have increasingly pushed the Earth towards an environmental breaking point. Even as researchers are racing to name and describe new species, they are simultaneously racing to save species from extinction. The spring after starting my backyard explorations, I discovered the existence of the Student Section at the nearby California Academy of Sciences in Golden Gate Park. The group offered activities after school and on weekends, along with occasional field trips to the One Green Earth Peter H. Raven RAVEN, P. H. 2020. ONE GREEN EARTH. ARNOLDIA, 78(1): 32-41 One Green Earth 33 surrounding countryside. The students also received a degree of access to the scientific departments at the academy, and by the time I was ten, I had become a regular visitor to the Entomology Department. There, I could compare and identify my beetle collections with the help of friendly curators, especially E. C. Van Dyke, a world expert on beetles who was always encouraging. By the time I was twelve, in the summer of 1948, I had begun to switch my interest to plants, largely because of a book called Manual of the Flowering Plants of California, by the great University of California botanist Willis Lynn Jepson. With the aid of this book, I could identify almost every plant species that I collected and determine whether there was anything unusual about the place I encountered it or the characteristics of the individual plants that I found. There had been no such book available for beetles. For plants, Jepson's Manual made the world seem small and knowable\u2014as if the different species in the Bay Area were parts of a large puzzle for me to discover and piece together. In the academy's Botany Department, curator John Thomas Howell (\"Tom\" to almost everyone who knew him) took me under his wing and taught me more each I time I visited him. I started helping in the department as a volunteer in 1948, and later that year, I was hired for my first job, sorting new collections that had come from people working with Tom around the state. My Early Exploration While I knew, even as a child, that botanists were still discovering new species of plants around the world, I had always assumed that plants in the region of California where I grew up were already well documented. Generations of earlier botanists had studied the flora, and it seemed as though all of the plants had already been named and included. My first personal experience with a new species began when I was in my final year at high school. Harlan and Margaret Lewis, who were preparing a monograph of the attractive native plant genus Clarkia, showed up at the academy. They were reviewing herbarium specimens of Clarkia, and they had come across an unusual one that I had collected a couple of years earlier on a slope of serpentine rock in the San Francisco Presidio. They wanted to grow it for their research, but it took me two years to find the colony again. When I finally sent them the seeds, they invited me to work with them at UCLA the following summer, between my junior and senior years at Berkeley. Following that experience, it was only natural for me to begin graduate work with Harlan in 1957. The unusual Clarkia eventually proved to be an unnamed species: Clarkia franciscana\u2014now a federally endangered species. While I maintained a lifelong interest in insects, I never looked back. At UCLA, I prepared a dissertation on a group of desert plants that were, like Clarkia, members of the evening primrose family, Onagraceae. At the age of twenty-two, I married a girl I had met at the student section, Sally Barrett, and the following year, somewhat to the consternation of my graduate advisors, we had our first baby. We had our second child, Elizabeth, in 1960, while we were living in London, where I had a postdoctoral fellowship at Kew Gardens and the London Museum of Natural History. We returned to California, and in 1962, after a job at Rancho Santa Ana Botanic Garden, I started what turned out to be a nine-year stint on the faculty at Stanford University. Fortunately for me, Stanford had a combined Department of Biological Sciences in which I had plenty of room to learn and grow in many aspects of the life sciences. Working with these colleagues, I could expand my research beyond its original emphasis on the classification of a particular group of plants and begin exploring topics with a broader and more theoretical footing. My closest colleague at Stanford was Paul Ehrlich, an entomologist and population biologist who has remained a mentor and friend for life. Comparing our thoughts on plants and butterflies, we recognized that the caterpillars of some groups of butterflies fed almost exclusively on one related group of plants. In these Facing page: A superbloom on the Carrizo Plain in California, with desert candle (Caulanthus inflatus) across the center of the image, the blue tansy phacelia (Phacelia tanacetifolia) in the foreground, and the hills beyond covered with hillside daisy (Monolopia lanceolata). PHOTOGRAPH BY ROB BADGER, FROM BADGER, R. AND WINTER, N. 2020. BEAUTY AND THE BEAST: CALIFORNIA WILDFLOWERS AND CLIMATE CHANGE. WINTERBADGER PRESS\/CALIFORNIA NATIVE PLANT SOCIETY. cases, few other kinds of butterflies fed on the same groups of plants. Cabbage butterflies, for instance, which I had observed in my childhood backyard, were among a group of related butterflies that fed on plants in the mustard and caper families. Paul and I came to understand that the ancestors of these plants had, over time, evolved chemical defenses that deterred most other insects. Ancestors of the cabbage butterflies, on the other hand, had gained the ability to break down or resist those defenses, which meant a whole food resource was more or less exclusively available to them. Paul and I developed, published, and named this stepwise process coevolution, which turned out to be one of the most fruitful scientific discoveries that either of us ever made. A couple of years earlier, my first Stanford graduate student, Dennis Breedlove had introduced me, through his fellow student and friend Brent Berlin, to a project that was being carried out in the Department of Anthropology. Professor A. Kimball Romney, one of the founders of cognitive anthropology and Berlin's graduate advisor, was working with colleagues to pursue various projects with the highland Mayans in the southernmost Mexican state of Chiapas. Together, the four of us conceived a project dealing with the names one group of these Mayans gave to the plants that grew in their area. Dennis moved to Chiapas for three years to carry out the botanical side of the study. We wanted to know what principles governed the way the Mayan community named their plants, Monarch butterflies (Danaus plexippus) advertise their poisonous nature by their bright colors and thus warn birds to leave them alone or suffer the consequences. This group of butterflies takes the process of coevolution one step further, getting poisons from the milkweeds on which their caterpillars feed and using them to protect themselves. HENRY DOMKE and what regularities we could find in comparing their system with those employed by groups from elsewhere. This Mayan community did not use a written language, and it turned out, this meant individuals only keep something like a few hundred plant names in their active memory. Within this system, they divided the kinds of plants most useful to them into many more categories than others. When this project began, the plants of southern Mexico were unfamiliar to me. Challenged with a rapidly growing number of herbarium cases filled with such plants, I had to find ways to name them in order to fulfill my part of the project. I eventually solved this problem with the help of many specialists, especially Jerzy Rzedowski, a Holocaust survivor who had become and has remained for many years the doyen of Mexican botanists, and the taxonomist Rogers McVaugh of the University of Michigan. On my next major collecting adventure in the tropics, where I served as an instructor for the Organization for Tropical Studies basic field course in Costa Rica during the summer of 1967, I was able to ship all the specimens to Bill Burger at the Field Museum in Chicago. He found a number of undescribed species among them and was quite pleased with what he received. Step by step, my interest in and knowledge about plants was expanding globally. Considering that my parents were living in Shanghai when I was born, and that my mother's grandfather arrived in California with his Irish family One Green Earth 35 Gunnera insignis was among the many wonderful new plants the author encountered while teaching a course for the Organization of Tropical Studies in Costa Rica in 1967. RICHARD MACK (Breen) as a member of the 1846 Donner Party, a global perspective had always been central to my family narrative. It seemed only natural that this perspective should be extended to plants. Research Coordination The mid to late 1960s were a tumultuous time for America, and all the more so for me. My wife, Sally, died of a sudden and unexpected health problem at age thirty. Our two young children were nine and seven at the time. This personal tragedy was compounded by the national unrest. These years were unsettling and extraordinary. This broader sentiment has been expressed well by Joan Didion, in her essay \"Slouching Towards Bethlehem,\" in which she describes the countercultural movement that had taken hold in San Francisco. \"Once we could see these children, we could no longer \u2026 pretend that society's atomization could be reversed,\" she concludes towards the end of the essay. \"This was not a traditional generational rebellion.\" This period saw the Tet Offensive, the assassinations of Robert Kennedy and Martin Luther King, the riots and arrests at the Democratic Convention in Chicago, and eventually the Kent State shootings. Demonstrations became an everyday event on the Stanford campus, as they did at other universities throughout the country. All in all, I became deeply confused about where the world was headed and uncertain about what the future held for me and, indeed, for the world. In this period, I worked with Helena Curtis, another biology writer, in preparing the first edition of what turned out to be a very successful botany text, The Biology of Plants. I also remarried relatively soon, to Tamra Engelhorn, whom I had met on the Organization for Tropical Studies course in 1967. Notwithstanding these positive events, I remained deeply troubled about the future and, indeed, about the purpose of life. My personal salvation came in the form of a sabbatical year in New Zealand, in 1969 and 1970. My intention in going there was to study the regional species of willow herbs, Epilobium, 36 Arnoldia 78\/1 \u2022 August 2020 Seiwa En, the Japanese Garden at the Missouri Botanical Garden, was designed as part of a plan to deepen community interest in this venerable institution (opened to the public in 1859) and, at the same time, to encourage an international outlook among St. Louisans. MARY LOU OLSON, MISSOURI BOTANICAL GARDEN the largest genus of the family Onagraceae. About a quarter of the roughly 160 species of the genus occurred in New Zealand and Australia, a strange fact considering the obvious New World origins of the family\u2014why were there so many species of Epilobium in that part of the Southern Hemisphere? They were all herbs but widely varied in appearance. They gave the impression of having evolved relatively recently and rapidly in the varied habitats of the region. As Tamra and I studied the Epilobium, we gradually regained our balance. New Zealand felt like a green paradise, and the people we worked with were level-headed, friendly, and helpful. One of them, Eric John Godley, the director of what was then the Botany Division at the Department of Scientific and Industrial Services, was of particular importance for me. We soon became fast friends. Nearly twenty years older than me, he calmly offered sound advice and joined us for enjoyable activities throughout our time in the country. The theory that the position of the continents had moved over geologic time had been proposed half a century earlier by the German geophysicist Adolf Wegener. His theory was essentially validated in the years just before we reached New Zealand, and it opened important new ways to interpret the origins of the plants and animals in the region. I was quick to apply them to the patterns about which I was learning and to publish the results. For example, the ridge that included New Caledonia and New Zealand separated from Australia and Antarctica (then still joined) about eighty-five million years ago, and most of the plants in New Zealand (including Epilobium) reached their new home by blowing or floating across the intervening seas. In later years, I presented similar interpretations in a series of papers with my geologist friend (and former member of my doctoral committee) Dan Axelrod. At our final dinner with Eric Godley, in the garden of his suburban home, he turned to me and asked what I was going to do next. He suggested that I might make the greatest contribution by emulating the great German botanist Adolph Engler, who, in the late nineteenth and early twentieth centuries, had led the production of the most important comprehensive works on plants, Die Pflanzenfamilien and Das Pflanzenreich. These works collectively described all the plants on Earth that were then known to science. Returning to Stanford, I kept Eric's advice in mind: I looked for a pathway to become a leader in encouraging others to undertake major projects, rather than simply continue to do my own research. The need for synthesis became obvious to me, and it has turned out over the years that instead of the roughly 250,000 species of vascular plants we had thought existed then, the actual number approaches twice as many. With these broader horizons in mind, I applied for the open position of director of the Missouri Botanical Garden in St. Louis during my single year back at Stanford. As matters turned out, I was successful. There I soon realized my efforts in coordinating and enabling the studies of others were more important than the results I could achieve as an individual, regardless of how useful and interesting the results of my efforts might prove to be. Global Collaboration The Missouri Botanical Garden is the oldest public garden in the United States. I had visited several times earlier to consult its excellent herbarium. On my arrival in 1971, the garden's only major research project, and the only one it had ever conducted abroad, was the Flora of Panama, which was being published serially as exploration and writing proceeded. My experience in Chiapas had taught me that to inventory the plants of a particular area properly it was necessary to live there and work with them daily. It seemed logical to find parts of the world that were of particular interest botanically, not being studied in detail by others, and to concentrate there. We began to hire staff with the help of several grants from the National Science Foundation and the support that accompanied the increased local interest in the garden that we were building. As the years went by, we were able to sponsor scientists to live in Nicaragua, Costa Rica, Ecuador, Peru, Bolivia, the Democratic Republic of the Congo, Madagascar, and Vietnam, and to form strong partnerships with a number of other countries. We established a branch office One Green Earth 37 for studying African plants in the herbarium at the Mus\u00e9um National d'Histoire Naturelle in Paris. Overall, these efforts led to the Flora Mesoamericana, a modern account of all plants between the Isthmus of Tehuantepec in southern Mexico and Panama; the revival of the Flora of North America, which covers the United States and Canada; national floras for many of the countries; and an online checklist of the plants of the Americas, a massive collaboration with institutions and researchers around the world that is headed by Carmen Ulloa Ulloa and Peter J\u00f8rgensen. I also helped to start and then coedit the Flora of China, a forty-nine-volume work that treats the more than thirty-two thousand species of plants found in the country where I had been born. This important and personally enjoyable project lasted for some three decades and was the product of a major cooperative effort between dozens of institutions and hundreds of individual botanists. It brought the botanists of China, taking up their new opportunities as the effects of the Cultural Revolution receded in the late 1970s, into cooperative contacts with botanists all over the world. The volumes were jointly published by Academic Press in Beijing and the Missouri Botanical Garden. Robert Woodson, the originator of the Flora of Panama, had told me in the course of a visit to St. Louis, back in 1961, that he thought they had accounted for nearly all of the species in the country. As our studies continued, however, we have reached the point where we now list approximately twice as many species from Panama as were known at that time. Everywhere botanists looked\u2014not only in Panama, but in North America, China, and around the world\u2014masses of new species turned up. Through this collective research effort, we were just beginning to grasp the magnitude of the Earth's biodiversity. The Need for Conservation When I was a young field biologist, in California, my studies were predicated on the assumption that the world would pretty much stay as it was. But in the 1960s, at Stanford, I started to 38 Arnoldia 78\/1 \u2022 August 2020 Libing Zhang (Missouri Botanical Garden), Hong Deyuan (Beijing Institute of Botany), Peter Raven, William McNamara (Quarryhill Botanical Garden), and Fu Chengxin (Zhejiang University), on the summit ridge of Huangshan (Yellow Mountain), in Anhui Province, China, on April 6, 2008. The field trip occurred a few days after a meeting of the Flora of China Editorial Committee, held at Zhejiang University. Hong and Raven were coeditors of the forty-nine-volume project. WILLIAM MCNAMARA become aware of the rapidly increasing destruction of nature around the world and the need to do something about it. California's population was then less than a quarter of its present 40 million, and the global population was less than a third of its present 7.8 billion, headed for 10 billion by the middle of the century. I began to worry about the severe effects of DDT and other pollutants\u2014Rachel Carson's Silent Spring appeared in 1962\u2014and I came to understand Paul and Anne Ehrlich's emphasis on population growth as a factor driving the destruction of ecosystems globally. In an effort to influence the 1968 presidential election, Paul and Anne published The Population Bomb, a real wakeup call about problems that were starting to become obvious. I was also becoming aware of the extensive destruction of tropical forests that was taking place; once I reached St. Louis this became a major issue for me in planning my personal activities and those of the garden. At the request and with the sponsorship of the National Science Foundation, I chaired a National Research Council study of priorities in systematic and evolutionary biology (Raven, 1974). By that time it had become obvious that the tropics were the most poorly known part of the Earth biologically and that the estimate for the number of species globally (then, with 1.5 million named, placed at 2 million) was much too low. Only five hundred thousand of the named species were tropical, yet two-thirds of the total number of species in well-known groups like plants and terrestrial vertebrates occurred there. It became clear that the actual number of species was at least 3 to 4 million\u2014and now we would probably say 20 million, with only 2 million of them yet named. In view of these new estimates and the fact that we had a sense that major habitat destruction was going on in the tropics, we selected focused research in the tropics as a top priority. We knew a lot less than we thought we did. Several years later, Bill Sievers, a program officer at the National Science Foundation, challenged me to head a study on setting specific research and conservation priorities in the One Green Earth 39 Tropical forests are being destroyed rapidly all over the world, as the 2011 clearing of primary forest for wood pulp in central Sumatra illustrates. WILLIAM LAURENCE tropics. I felt that we first needed a more comprehensive understanding of the degree and rate of destruction that was going on in the tropics. This information would help us set the most critical priorities during our study. It seemed to me that the man for the job was Norman Myers, an imaginative English ecologist and conservationist. He had gone to Kenya in the British Colonial Service and stayed on after independence, working as a teacher, guide, photographer, and consultant. By the late 1960s, Norman had become one of the very first to recognize that we were entering a major extinction event and to write about it. To conduct the study of tropical forest destruction for us over an eighteen-month period, he visited all of the major tropical areas and many tropical forested countries, consulting a great deal of \"gray literature\" and conducting interviews. His report, published in 1980, proved to be a bombshell, documenting rates of deforestation much higher than were generally assumed at the time. It spurred us all to higher levels of action, given the urgency of the task facing us. Once these relationships had become clear to me, I decided to devote a large part of my energy and available time to accomplish what could still be done while our present wealth of organisms and their ecosystems still exist. I had the opportunity to present the case for action at the American Association for the Advancement of Science annual meeting in Chicago in 1987. It was a large audience, and many people told me later that they had heard about the problem of mass extinction for the first time then (Raven, 1987). Even at the lower estimates of species numbers with which we began, we learned that for every twenty species of plants, animals, and other organisms in a given forest, nineteen were still unknown. So when an area of tropical forest is cleared, the overwhelming majority of species were disappearing without being documented by scientists. The problem was becoming generally obvious. Today, more than a quarter of the tropical forests standing when the Convention on Biological Diversity was ratified, twenty-seven years ago, have been cleared. The rate is only increasing. Few researchers project that any substantial stands of tropical forest will remain by the end of this century. At the same time, the world climate is warming rapidly, with no strong international agreement in place to slow it down. Biologists can still hope to fill out a relatively complete picture of species numbers and distribution for vascular plants, terrestrial vertebrates, and a few other groups of organisms. But carefully constructed sampling protocols afford the only hope for learning much about groups such as nematodes, mites, and fungi, for which we have recognized fewer than one in a hundred species yet. At least a quarter of all species, most of them unknown, are predicted to disappear from the face of the Earth by the close of this century. What we find and save now will be all we can pass on to those who come after us. We have a moral obligation to do so. As University of Pennsylvania biologist Daniel Janzen has remarked, \"If we don't save it now, we can't save it later.\" Collective and Individual Action When I was lying in bed recovering from measles at the age of seven, nearly eighty years ago, it would have been impossible for anyone, and certainly for me, to imagine the tremendously difficult problems we are facing now. Estimates by Global Footprint Network, based on United Nations statistics, reveal that human demand on natural resources in 1961 corresponded to about 73 percent of what Earth could renew at the time. Our demand has risen to 175 percent currently (Lin et al., 2018; Global Footprint Network, 2019). In other words, by July 29, 2019, humans had demanded as much of the Earth's resources as those ecosystems could regenerate in the entire year. Taking this depletion on a per-person basis, we find the averages in the United States, Gulf Countries, and Western Europe are the highest. In contrast, the averages within countries that lack ecological resources and purchasing power reflect very low demands, indicating extreme deprivation and difficult material prospects for their residents (Wackernagel et al., 2019). Huge inequities also exist within nations. Schemes for conservation imposed by wealthier nations tend to be massively unjust towards poorer nations, which have far fewer resources to devote to them than their wealthy counterparts. If the richer nations 40 Arnoldia 78\/1 \u2022 August 2020 would partner with them and help financially, the schemes could work, but there is little sign of such mutual respect and the love that it would require to generate such help. During my career, I have become convinced that only global collaboration and understanding can give us hope for sustainable life on Earth. Any such collaboration must be based on social justice and a spirit of love and understanding between people everywhere. Yet global success ultimately requires individual action, and it can exist only in a socially just world. Each of us must learn as much about the world, and especially about the poorer parts of the world, as we possibly can. We must live as sustainably as possible. We must vote for politicians who try to understand what's going on beyond their own short terms of office and who recognize the critical importance of arresting and then reversing global climate change. We must support the preservation of the species and ecosystems living today. We must also find ways to gradually lower our population to a level that the planet can support, instead of continuing to pretend that our global resource consumption doesn't matter. All of these actions are predicated on a fundamental need for us to find ways to love and appreciate one another. Our civilization is very young and vulnerable. Our ingrained habits of selfishness and competition were doubtless beneficial in a world where the total human population numbered in the hundreds of thousands, but they have become a sure pathway to destruction now. It is clearly time for us to act. Acknowledgments I sincerely thank Mathis Wackernagel, Global Footprint Network, for his suggestions, and acknowledge the exceptionally fine work of Jonathan Damery, the editor. Literature Cited Global Footprint Network. 2019. 2019 edition of the National Footprint and Biocapacity Accounts. Retrieved from https:\/\/data. footprintnetwork.org. Lin, D., Hanscom, L., Murthy, A., Galli, A., Evans, M., Neill, E., Mancini, M. S., Martindill, J., Medouar, F.-Z., Huang, S., et al. 2018. Ecological footprint accounting for countries: Updates and results of the National Footprint Accounts, 2012-2018. Resources, 7(3): 58. Retrieved from https:\/\/ www.mdpi.com\/2079-9276\/7\/3\/58. Myers, N. 1980. Conversion of tropical moist forests. Washington, DC: United States National Academy of Sciences. Raven, P. H. 1974. Trends, priorities, and needs in systematic and evolutionary biology. Brittonia, 26: 421-444. Raven, P. H. 1980. Research priorities in tropical biology. (P. H. Raven, Chairman, Committee on Research Priorities in Tropical Biology, National Research Council.) Washington, DC: United States National Academy of Sciences. Raven, P. H. 1987. We're killing our world: The global ecosystem in crisis. MacArthur Foundation Occasional Paper; reprinted in Missouri Botanical Garden Bulletin, 75: 3-7. Wackernagel, M., Lin, D., Evans, M., Hanscom, L., and P. H. Raven. 2019. Defying the Footprint oracle: Implications of country resource trends. Sustainability, 11(7): 2164. https:\/\/doi. org\/10.3390\/su11072164 Peter H. Raven is president emeritus of Missouri Botanical Garden, in St. Louis. In 2014, he received an honorary doctor of science degree from Harvard University. His autobiography, Driven by Nature: A Personal Journey from Shanghai to Botany and Global Sustainability, will be published by the Missouri Botanical Garden early in 2021. One Green Earth 41 Pat and Peter Raven by the side of the rapidly receding Portage Glacier in Alaska, on a trip with scientists and Evangelical Christians to study climate change in early May 2007. PHOTOGRAPH BY CAMILLE PARMESAN, EDITED BY HENRY DOMKE"},{"has_event_date":0,"type":"arnoldia","title":"Each Year in the Forest: Summer","article_sequence":5,"start_page":42,"end_page":51,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25712","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24eab27.jpg","volume":78,"issue_number":1,"year":2020,"series":null,"season":null,"authors":"Hipp, Andrew L.","article_content":"Each Year in the Forest: Summer Andrew L. Hipp Illustrated by Rachel D. Davis HIPP, A. L. AND DAVIS, R. D. 2020. EACH YEAR IN THE FOREST: SUMMER. ARNOLDIA, 78(1): 42-51 I Each year, for a few weeks in succession, I tell myself that we are done with spring. I think the chorus frogs have stopped singing for the year. Then they start up again. The sedges bloom in a flurry, anthers waving, and they die back. But then another wave arrives. Warblers stream through town and then are quiet. A few days later, I hear that a blue-winged warbler was lurking in the western suburbs. There is no clean joint between spring and summer. Still, near the beginning of June, the great waterleaf plants that emerged as blotchy, hairy leaves in early April become splendid with gauzy blue flowers. Dagger-like fruit columns sprout from the center of wild geranium flowers, arising from between the nutlets that develop at the bases of the petals. After a few weeks, they will catapult their seeds into the woods around them, then persist, irrelevant as the flowers fall to pieces. Sugar maple leaves unfurl. Toothwort begins yellowing; false mermaid becomes flattened and desiccated; the culms of straight-styled wood sedge flatten out as though they'd been stepped on by elephants. Leaves of wild leeks do the same. The forest prepares itself for the long run uphill to the solstice and over the crest to autumn. Bur Oak Acorns Forest: Summer 43 In early June, hackberries and basswood leaves are still expanding. Oaks spill into flower as gray squirrels nip off the shoot tips and strew them across the forest floor. The door continues to close on spring for a few more weeks: Virginia waterleaf flowers nod at the bases of white oaks; great crested flycatchers and red-eyed vireos call at intermittent points along every walk in the woods; sedges of the second flush drop their fruits (nutlets wrapped in papery green perigynia that will see them through to fall), and the sedges of the third flush begin to ripen; woodland bluegrass spikelets gush with anthers. But spring isn't over until the last leaves are fully open and the holes close in the canopy. At that point, the forest floor becomes its darkest, and leaves of the white bear sedge abruptly broaden out. Summer spreads out before us with its long weeks of photosynthesis in full spate. Its progression of insect songs will take us through hot afternoons and humid mornings to the end of September. Fireflies make their first flights near the beginning of June without any obvious chance to warm up and no training: they've only just pupated and emerged as adults. Still, they seem not to worry that their prospective mates might notice how raw and unpracticed they are. After a season of solitude as infants, after never having had a mating season at all, they mature, illuminate, and fly. They are nonchalant. They exhibit none of the uncertainty of young vertebrates, the stumbling fawns and warbling young white-throated sparrows, the clumsy great-horned owl babies. Instead, male fireflies execute aerial wooing dances whose precise choreography they are born knowing. These flights differentiate them by species. One flies a J-stroke. Another flies in dots and dashes. Meanwhile, the females sit in nearby shrubs and blink, drawing the males in.1 I was once swinging a double-bladed weed cutter through the University of Wisconsin's Curtis Prairie near sunset and found that I had attracted the attention of a firefly, who flew in toward me and tried to attract the fancy of the bolts glinting along the joint between the handle and the blade. It was unsuccessful. Mosquitoes are drawing blood by this time, bringing common nighthawks out into the open. I notice the nighthawks in the evenings when their splattery \"peent,\" juicier and less strident than the woodcock's, draws my attention to the sky. They career around overhead with an agility that defies physics, banking almost as sharply as dragonflies, diving and pulling up, the white bands on their wings flashing. All the while, they are funneling insects into their mouths. Watching them fly, I cannot understand how a diet of small insects can sustain such energy. Their flight is ceaseless but for the rare times that I have seen one perching, roughly the size and shape of a large bread loaf, on the edge of a flat city rooftop. I sometimes hear a male roaring as he drops through the sky to impress a potential mate, wind booming through the primary feathers as he nears the bottom of his dive. This is the same time of year when I used to hear a close relative, the whip-poor-will, calling in wooded neighborhoods of the University of Wisconsin-Madison Arboretum on my late-night bike rides home from work. 44 Arnoldia 78\/1 \u2022 August 2020 All through the month, I watch the juneberries slowly ripen. Many people grow them in their front yards and apparently have no idea that they are edible. When the berries finally become dark and soft and sweet enough to eat, they are delicious, and for a couple of weeks each June, I cannot walk past the vacant lot turned playground down our street without standing at the fence for at least a minute or two, eating berry after berry. They do not travel or store well, so I eat as much as I can while they are in season. As I do, I know without a doubt that we are in the midst of summer, though I couldn't tell you exactly when it started. II In the weeks leading up to the solstice, the great waterleaf petals fall, and their spidery inflorescences bear capsules that ooze pulpy white ovules when you pinch them. Jewelweed grows to chest height and produces mysterious translucent fruits that appear to have preceded the flowers. They are, in fact, the products of cleistogams, flowers that never open, in which the stigmas and anthers are closed in together and external pollination is excluded. They are the plant's answer to the risk of not getting pollinated. They explode in hand before I notice a single outcrossing flower. Beside them, leaves of the wood nettle sprout translucent galls that resemble tapioca pearls with a dark core. Inside each grows a gall midge, Dasineura investita, which I have probably seen in adulthood but never recognized. Wild leek scapes poke up through the leaf litter, each tipped with an arrowhead-shaped hood. Some colonies emerge erect, others arched over and darkened on all surfaces. The flowering stalks sparsely map the extent of the dense swards of green foliage that grew fast in March and April, dissolving into the soil a couple of weeks before the scapes emerged, as the last holes closed in the canopy. There is no evidence of the leaves as the inflorescences swell against the hoods and tear through their sides. The inflorescences stand for a few days like fists raised above the leaf litter. The flowers open gradually, six papery tepals spreading beneath a congregation of stamens. Jewelweed Wild Garlic Forest: Summer 45 The individual flower stalks grow as the flowers continue to open, so that at their peak, the colony of leeks is a cloud of airy white inflorescences. Morbid owlet moths waft across the path and settle onto the undersides of last year's sugar maple leaves. The moths are similar to the faded maple leaves in color and value. Leconte's haploa moths flap their black-striped white wings like flags and settle on the shorter plants. Ebony jewelwing damselflies bumble along near creeks and at the edges of woodland marshes. They are so faithful to their habitat that when I see one, I know\u2014whether I am in central Wisconsin, northern Illinois, or overseas2\u2014that I am near water. They have a flair that I appreciate and a casual gait that I admire, flapping and gliding, bouncing between plants rather than vigorously taking out insects over open water. They seem to lack the ruthless efficiency of the other damselflies and dragonflies in the neighborhood. Summer pricks the forest floor with light. The delicate white petals of enchanter's nightshade open at the tips of bristling ovaries. Wild garlic spathes open to reveal a cluster of bulbils atop a thin scape; flowers emerge from among the bulbs and turn toward the canopy on narrow stalks. Jumpseed that started the year as red-stained leaves low to the floor produce flowers roughly the size and shape of cooked orzo. Honewort flowers wink on like stars scattered between the major constellations and then give way to plump, rubbery fruits that smell of celery leaves. Ripened spikelets of fowl mannagrass shine at the tips of slender branches; I gather a handful and drizzle them along the side of the path, where they crackle like grains of couscous against the leaves. Seeds ripen to a glossy chestnut brown inside the wild ginger berries lying in the duff, each rimmed with an oily crest, an elaiosome, that is as delicious to ants as a fresh-baked roll is to a human. A dull, hairy capsule of the great waterleaf is nestled within the persistent calyx. But when I slough off the fruit wall with my fingers, the seeds inside lie wet and pearly white, reflecting sunflecks. Then we hit the solstice. There is no more to see on this day than on any other, but we have a few extra minutes in which to see it.3 For a month or Ghost Pipes Morbid Owlet Moth 46 Arnoldia 78\/1 \u2022 August 2020 so, the woods will trundle, pause, grow, and decompose. The long days will fill with cicada songs and greenery, the nights with clouds of fireflies. It is downhill now in both directions. III Through June and early July, the fruiting bodies of dead man's fingers (Xylaria) 4 curl from gaps in fallen tree trunks and protrude from the chipped trails. They are powdery gray and tipped with white at first, blackening as they mature. Collared parachute mushrooms (Marasmius rotula) sprout from decomposing branches and tree fragments. Their caps are gelatinous and crenate. Their ridged margins droop down over the tops of the stems like children's umbrellas that leave only torsos and legs visible as they walk the rainy paths. Orange mycena mushrooms (Mycena leaiana) sprout small colonies along the sides of rotting red oak trees. Trooping crumble caps (Coprinellus disseminatus) mass up in the wreckage of fallen branches, fragile and diminutive forests forming in the canopy's cemetery. White jelly fungus (Ductifera pululahuana) glistens at knee level. Ghost pipes emerge from the forest floor, white, with nodding flowers that look down toward their toes, feeding off the fungi that live with the tree roots. The forest is growing at full tilt, and already it is being devoured. In Maple Grove Forest Preserve, there is a brown, spongy, decaying white ash trunk5 that I regularly check for fungi. I found it bristling, on the Fourth of July last year, with what I thought were bright red, tiny toadstools. The caps were less than a millimeter in diameter, the stalks threadlike. A flat-backed millipede was crawling among this fur of filaments and pinheads. On the side of the log, a white, fleshy Crepidotus, a common woodeating fungus, had emerged with a cluster of something that resembled tiny puffballs, a bit larger than mung beans. On a nearby log, the Xylaria were blackening at the tips. I posted the \"toadstool\" photos to iNaturalist, a social networking site for sharing and discussing biodiversity photos. Within hours, I had heard from a Tasmanian naturalist6 who observed that while the Crepidotus and Xylaria were fungi, the others were not. They were slime molds: Cribraria for the Dead Man's Fingers Orange Mycena Forest: Summer 47 \"toadstools,\" wolf's milk (Lycogala epidendrum) for the \"puffballs.\" I was surprised and delighted. In an afternoon, I'd found a whole new branch of the tree of life to watch for in the woods. I looked up the Encyclopedia Brittanica entry on slime molds:7 Science fiction did not invent the slime molds, but it has borrowed from them in using the idea of sheets of liquid, flowing protoplasm, giant voracious amoebae, engulfing and dissolving every living thing they touch. What fiction could only imagine, nature has evolved, and only their sharp dependence on coolness, moisture and darkness has kept the slime molds from ordinary observation, for they are common enough. Two days later, I returned to the woods to find that I could now distinguish at least six slime molds by eye. In addition to Cribraria and wolf's milk, I found Arcyria cinerea, which looked like grains of rice suspended by threads; Tubifera, pincushions on the sides of the logs; an undifferentiated yellow plasmodium that might have been Physarum crawling over the surface of the log; and the aptly named dog-vomit slime mold (Fuligo septica) mounded up on logs stripped of their bark. I slapped the colony of Cribraria: spores rose and formed a fog around my hand, then drifted off along the length of the log. After the slime mold sporing bodies disintegrated, the tree lay more or less naked. I have watched one end of the trunk crumble over the past year, trampled to the point that it now grades into the trail. The slime molds themselves didn't do this: they live largely on bacteria and fungi, not wood.8 Ants have trailed through the sapwood and replaced xylem with frass; mycelia of chicken of the woods (Laetiporus sulphureus) have wound through the tree, devouring lignin, leaving the wood blocky and red; the roots of jewelweed and enchanter's nightshade growing on the top have tunneled into the wood; moss growing on the shady side has helped keep the tree spongy. Slime molds help with the mop-up. When I returned this year, small patches of honeycomb coral slime mold (Ceratiomyxa fruticulosa) appeared on the side of the log by the first of July, along with wolf's milk. Then, on the third of July, as though on cue, a coat of Cribraria sporangia appeared on the flattened rubble pile along the path. IV By late July, many plants are scarred by insects. On the midribs of jewelweed leaves, translucent swellings conceal young gall midges (Neolasioptera impatientifolia), which grow as tiny yellow larvae inside each blister.9 Elm-leaved and zigzag goldenrod that haven't come into flower yet are inscribed with meandering leaf-miner tunnels that begin small, thicken as the larvae inside grow, and often terminate in a hole. Leaf miners find white snakeroot as well. Other leaves are crisscrossed with slime trails that I suspect are left by slugs or snails, but the leaves often show no evidence of chewing or scraping damage. The broad, soon-to-become-evergreen leaves of white bear sedge begin to resemble subway maps, with routes scratched into the mesophyll by leaf-miner flies (Cerodontha sp.), who follow the veins of the leaves longitudinally, tunneling in parallel before they veer diagonally to connect the paths. Along a trail through the Morton Arboretum's East Woods, woodland sunflowers are packed as densely as a planted field. They flower in July, their brilliant yellow faces all turned intently southward, extending almost as far as I can see in the shade of the white oaks. Tall bellflower comes into bloom one day, and the blue flowers are high enough to stare me in the eye, a single style snaking out from the white-target center of each flower. Shining bedstraw scrambles along like baby's breath at ankle-height. False nettle erects columnar inflorescences that angle from the leaf nodes and look strong enough to hang a coat on. The filigree of wood nettle inflorescences signifies the end of one's opportunity to harvest the leaves for the year. Before this, they can be boiled and eaten, though in my experience they are bland. After this, I have been told, they become bitter. Perhaps this marks the beginning of the end of summer. The interval from mid-July to the middle of August is hot and slow. I lose track of what is going on in the woods. I travel for a conference and come back to find that Solomon's seal berries are ripe; when I last looked, in early July, they had just broken out of the papery corollas that enclosed them. We leave on vacation, north to where Canada mayflower is in fruit and clubmosses are thick in the shady portions of the forest, and we return to find bottlebrush grass looking ragged, wild leeks beginning to fruit, clearweed in flower. Moonseed sprawls over fallen logs. This is the last month of summer before the boys return to school. The days spread out like a fog low over the field in the early morning, amorphous, hot, hard to pin down. V As summer draws to an end, sounds of the fields, woods, and suburbs mark my progression through each day. Between mid-July and early August, the robins relinquish their predawn singing to the cardinals. This changing of the guard always catches me unawares. Crickets stop singing as the sun starts to bear down. Cicadas and lawnmowers fill the midafternoon. Our family bustles around with supper, kitchen noises spilling out into the yard, then robins begin chuckling in the neighborhood. Cicadas give way to crickets about thirty or forty minutes after the sun sets. Crickets sing through the screen well into night as I sit by the window or by the fire in the backyard. White snakeroot comes into bloom along trails and on the margins of woods by the baseball fields and parking lots. White baneberry fruits ripen, and the stigmas, shriveled at the tip, form a black eyeball. Inside are half a dozen glistening, wedge-shaped brown seeds embedded in pulp. Wingstem blossoms in the floodplains beneath the silver maples. Black elderberry ripens to sprays of small, dark berries. American pokeweed berries swell green and darken along one edge, filled with black, lenticular seeds. The brittle, 48 Arnoldia 78\/1 \u2022 August 2020 Forest: Summer 49 jumpy, clingy fruits of the pathway species ripen. Lopseed fruits become completely reflexed. Jumpseed flowers enclose brittle fruits that have been hardening over the previous weeks and now spring at a touch. Fruits on enchanter's nightshade become bristly. Stickseed transitions from immaculate white flowers\u2014five petals, no longer than a millimeter or so, each encircling a donut of tissue (the fornices) that extends up from the flower's throat\u2014to stick-tights that will give you hours of work if you brush against a single plant while wearing a sweater. Last year, red oak acorns littered the trails by the middle of August. I worried that they were falling too early, that they were all rotten, but biting a few open, I found mostly healthy cotyledons filling the shell. I floated out a sample at home, and about 50 percent sank, suggesting they were viable. But these were still at the outset of their journey, and not all would survive on the forest floor. Within hours of landing, they might be visited by insects who tunnel in and devour and fill the shell with frass before they depart, poor house guests. They might be preyed upon by molds and other fungi. The bur oak acorns swell through August and begin falling near its end, caps clothed in a ruff of kinked scales. Like the red oaks, they are in danger as soon as they land: pop the caps off of fallen bur oak acorns, and you often find writhing yellow larvae dying to get into the meat of the nut. The white oaks and Hill's oaks generally hang on a bit longer before they release their progeny to the ravaging insects, fungi, squirrels, and jays. Oak leaves and stems balloon up with galls of all types. By the end of the month, katydids rasp from the treetops. There are no sutures between the seasons. We can flip over every log and scrabble around on the forest floor, and we'll find a multitude of signposts: false mermaid seedlings firing up, proliferations of mycorrhizae, cicadas Woodland Sunflower White Baneberry emerging, earthworms growing torpid as temperatures drop. With so much to choose from, we might as well start the forest year here, with the red oak acorns raining down to their various fates. As they bed down and some, at least, find a safe place to get a radicle into the ground, they are staking out a part of the forest that they may work for centuries. They have as strong a claim on the beginning of the year as anyone does. Endnotes 1 For a wonderful discussion of firefly biology and the importance of the dances to firefly taxonomy, read, Evans, H. E. 1968. In defense of magic: The story of fireflies. In Life on a little known planet: A biologist's view of insects and their world (chapter 6). New York: Dutton. 2 In spring 2014, living west of Bordeaux with my family, my commute to work often included a bike ride from the train station at Gazinet through a sandy, spring-fed forest with a little creek. One morning, near the beginning of summer, I spotted what I thought to be the ebony jewelwing I had learned on the Lower Wisconsin River, and I saw it several times more during the last weeks of our stay. It turned out, though, that the species I knew, Calypteryx maculata, is endemic to Eastern North America. But the genus has Eurasian relatives as well. It seems most likely I was following the beautiful demoiselle, Calypteryx virgo, which lives along fast-flowing streams across much of Europe. 3 Tim Dee writes of the day after the winter solstice: \"The extra minute [per day] had nothing more to show than what was already present - it showed just a minute more of that. More light but, so, all begins again. Today, there was nothing else to see but there was one more minute to see it in.\" Dee, T. 2020. Greenery: Journeys in springtime. London: Jonathan Cape. 4 I use the common name loosely here to refer to the fungal genus Xylaria, whose species are not easily distinguished from one another without microscopic study that I have not undertaken. 5 I am indebted to my colleagues Christy Rollinson and Ross Alexander (at the Morton Arboretum) for their help identifying this tree from a wood sample. 6 Lloyd, S. 2019. Tasmanian myxomycetes. https:\/\/sarahlloydmyxos.wordpress.com\/ Red Oak Acorn 50 Arnoldia 78\/1 \u2022 August 2020 PLANTS REFERENCED Forest: Summer 51 Acer saccharinum - silver maple Acer saccharum - sugar maple Actaea pachypoda - white baneberry Ageratina altissima - white snakeroot Allium canadense - wild garlic Allium tricoccum - wild leek Amelanchier sp. - juneberry Asarum canadense - wild ginger Boehmeria cylindrica - false nettle Campanulastrum americanum - tall bellflower Cardamine concatenata - toothwort Carex sp. - sedge; there are other sedge genera, but these are the \"true sedges\" that dominate in our woodlands Carex albursina - white bear sedge Carex radiata - straight-styled wood sedge Celtis occidentalis - hackberry Circaea canadensis - enchanter's nightshade Cryptotaenia canadensis - honewort Floerkea proserpinacoides - false mermaid Fraxinus americana - white ash Galium concinnum - shining bedstraw Geranium maculatum - wild geranium Glyceria striata - fowl mannagrass Hackelia virginiana - stickseed Helianthus strumosus - woodland sunflower; though the colony I am referencing in the Morton Arboretum's East Woods may be referable to H. decapetalus Hydrophyllum appendiculatum - great waterleaf Hydrophyllum virginianum - Virginia waterleaf Hystrix patula - bottlebrush grass Impatiens sp. - jewelweed, touch-me-not Laportea canadensis - wood nettle Lycopodiaceae - clubmosses (various genera) Maianthemum canadense - Canada mayflower Menispermum canadense - moonseed Monotropa uniflora - ghost pipe Persicaria virginiana - jumpseed Phryma leptostachya - lopseed Phytolacca americana - American pokeweed Pilea pumila - clearweed Poa sylvatica - woodland bluegrass Polygonatum biflorum - Solomon's seal Quercus alba - white oak Quercus ellipsoidalis - Hill's oak Quercus macrocarpa - bur oak Quercus rubra - red oak Sambucus canadensis - black elderberry Solidago flexicaulis - zigzag goldenrod Solidago ulmifolia - elm-leaved goldenrod Tilia americana - American basswood Verbesina alternifolia - wingstem Andrew Hipp is the senior scientist in plant systematics and herbarium director at the Morton Arboretum in Lisle, Illinois. He conducts research on the origins and implications of plant diversity, with a focus on oaks, sedges, phylogenetic ecology, and trait evolution. You can read about his research at http:\/\/systematics.mortonarb.org and follow his natural history blog at https:\/\/botanistsfieldnotes.com. Rachel Davis is an independent visual artist in the Chicago area. She works at the interface of natural science, abstract painting, printmaking, and textiles, integrating the formal and empirical elements of the natural world in her work. You can see more of her work at https:\/\/artbumble.com and follow her on Instagram: @art_bumble. 7 Cohen, A. L. R. 1969. Slime molds (slime fungi). In: Encyclopaedia Brittanica (Vol. 20). Chicago: William Benton. 8 Stephenson, S. L. and Stempen, H. 1994. Myxomycetes: A handbook of slime molds. Portland: Timber Press. 9 For an image of the gall and the larva inside: Hatfield, M. J. 2013. Cecidomyiidae, jewel weed gall - Neolasioptera impatientifolia. Bug Guide. https:\/\/bugguide.net\/node\/ view\/741909"},{"has_event_date":0,"type":"arnoldia","title":"Speak, Cottonwoods","article_sequence":6,"start_page":52,"end_page":53,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25713","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d24eab6b.jpg","volume":78,"issue_number":1,"year":2020,"series":null,"season":null,"authors":"Wheeler, Emily","article_content":"In August 1895 and September 1896, the dendrologist John George Jack visited his hometown of Chateauguay, on the shores of the St. Lawrence River, near Montreal. By then, Jack had been working at the Arnold Arboretum for a decade. Director Charles Sprague Sargent had hired Jack as a manual laborer in 1886. The self-educated Jack rose over time to become a popular lecturer, a prolific plant collector, and an associate professor of dendrology at Harvard. Jack returned to Boston from Chateauguay with seedlings of the eastern cottonwood (Populus deltoides). Fifteen members of accession 16611 were planted out near Peters Hill. In 1894, Jack had written a five-part series about the trees and shrubs growing near Montreal in the magazine Garden and Forest, which Sargent oversaw. In the final section, Jack described an impressive cottonwood growing near the Chateauguay River: \"This tree is said to have been not much more than a sapling within the memory of some of the older inhabitants.\" Already, in 1894, its trunk measured more than five feet in diameter. This remarkable tree bore only pollen- producing male flowers (cottonwoods are dioecious), but the seedlings Jack collected may well have come from nearby. The seed capsules ripen in early summer and burst to release tiny seeds attached to cotton-like strands\u2014nature's dust bunnies. A single tree can produce up to forty million seeds. According to the Arboretum's records, Jack made plant collections near Chateauguay on a near-annual basis through 1912 (the year his mother, Annie, a well-known horticultural writer, died). Cottonwoods were a repeat interest. Another singular tree grew on an island in the mouth of the Chateauguay River, where a convent was then located. Jack had collected an herbarium specimen from this tree in 1889, and he would collect additional specimens from the same tree on at least four other occasions. The undersides of the leaves were more silvery than those of other cottonwoods, and Sargent came to recognize the tree as a hybrid between the eastern cottonwood and the balsam poplar (Populus balsamifera). In 1913, he named the hybrid in Jack's honor: P. x jackii. Although Jack collected cuttings from the hybrid, none of the resulting trees are living today. As for the original accession of eastern cottonwoods, five remain, all at the juncture of the Peters Hill loop and the short oak-lined spur leading to Poplar Gate. These are large trees. Their silvery gray bark has matured into deep furrows. The trunk diameter of the largest is around three feet, which is impressive but not nearly the five-foot specimen that Jack had observed towering over the rich bottomlands. In 1950, a year after Jack died at his farm in Walpole, Massachusetts, his name would appear in the New Yorker, in an essay by Vladimir Nabokov. (The essay was later adapted as the final chapter of Nabokov's memoir, Speak, Memory.) \"I would like to have the ability Professor Jack, of Harvard and the Arnold Arboretum, told his students he had\u2014of identifying twigs with his eyes shut, merely from the sound of their swish through the air,\" Nabokov wrote. Nabokov had settled in the Boston area in 1941. Although he was an academic with a literature degree from Cambridge, England, he had, from boyhood on, a deep love of nature\u2014 especially butterflies. (He even worked with the lepidopteran collections at Harvard's Museum of Comparative Zoology.) Whether Jack and Nabokov met is unknown. Yet the writer spelled out examples of trees Jack's auditory keenness could identify: \"hornbeam, honeysuckle, Lombardy poplar.\" Leaves of the eastern cottonwood, like those of the Lombardy poplar (Populus nigra), have flattened petioles. Even the faintest breeze can cause the leaves to rustle. To some, the leaves of the eastern cottonwood shiver. If it's true that Jack could identify plants by their sound, perhaps, in the case of poplar, he was remembering the sound of wind caressing the big leaves of a mature cottonwood grove along the Saint Lawrence River\u2014perhaps remembering the trees that drew him back again and again. Emily Wheeler lives in Jamaica Plain and is a docent at the Arboretum. Speak, Cottonwoods Emily Wheeler WHEELER, E. 2020. SPEAK, COTTONWOODS. ARNOLDIA, 78(1): 52-53"},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25695","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270a36e.jpg","title":"2020-78-1","volume":78,"issue_number":1,"year":2020,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Uncommon Gardens","article_sequence":1,"start_page":2,"end_page":5,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25700","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270af6b.jpg","volume":77,"issue_number":4,"year":2020,"series":null,"season":null,"authors":"Goulet-Scott, Ben","article_content":"With one last gulp of iced tea, I stepped out of a rented sedan onto the weedy shoulder of Forest Service Road 117 to perform my pre-fieldwork ritual. I tucked my pants into my socks, applied sunscreen and bug spray, and pressed a baseball cap over my spiky bed head. The morning temperature in western Kentucky was already approaching 90\u00b0F (32\u00b0C), unusual for late April. I grabbed my water bottle and tablet from the back seat and turned towards my experimental garden plot, which was planted with three subspecies of pink-flowered herbaceous Phlox. As a doctoral student working with Robin Hopkins, a faculty member at the Arnold Arboretum, I have returned regularly to western Kentucky and Tennessee to study the role of local adaptation in the divergence and speciation of these closely related lineages. A skeletal dead tree stood on the opposite side of the field, a favorite perch for large birds. I recognized the familiar broad-shouldered silhouette and gleaming white head of an adult bald eagle. Surely it had long since noticed me and my car, and as I pushed through the tall Uncommon Gardens Ben Goulet-Scott grass to arrive at my modest garden, I wondered if I might now be familiar to the eagles of this area. I was relieved to find that my plants still stood, and in fact, they seemed to be thriving. The spring before, in 2018, I had worked with Robin and lab technician Matt Farnitano to plant 321 rooted cuttings at this site, each no more than four inches tall. Now many of the plants boasted dozens, even hundreds, of bright pink flowers. I set down my water bottle and turned on the tablet, ready to record herbivore damage and count flowers for as long as the daylight permitted. This plot is a type of experiment known as a common garden. Three different taxa\u2014Phlox pilosa subsp. pilosa, P. pilosa subsp. deamii, and P. amoena\u2014had been planted in a random order, and because the growing conditions are consistent, any differences in traits among the three taxa can be ascribed to genetic differences rather than plastic responses to the environment. Common garden experiments have a rich history in plant biology. Botanists in the first half of the twentieth century (especially G\u00f6te Turesson, Jens Clausen, David D. Keck, and Phlox Common Garden 3 Facing page: Intermixed Phlox subspecies flower in the author's common garden in the Land Between the Lakes region of western Kentucky. ALL PHOTOS BY THE AUTHOR William Hiesey) made foundational contributions to our current understanding of heritable variation in natural populations using common gardens. Outside the Weld Hill Research Building at the Arnold Arboretum, other researchers are using a series of common garden plots to study the ecology, morphology, and physiology of woody plants. In fact, the entire Arnold Arboretum can be viewed as a large common garden, with plant species and varieties from around the world growing in one location. My research in Kentucky required not one but three common gardens, one in each habitat of my three study taxa. During the summer of 2017, I had traveled throughout the native ranges of these three subspecies in the southeastern United States and collected plant material for the gardens. Perennial Phlox propagate well from cuttings, so I collected single stems from wild plants, leaving the rest of the plant in the ground. I mailed these stems back to labmates at the Weld Hill Research Building who planted them in soil so they would produce roots. After one year in the Weld Hill greenhouses, they furnished three cuttings each, allowing me to plant a genetically identical panel of cuttings in each garden. All three of my common gardens sit adjacent to a wild population of one of the three subspecies. This experimental design\u2014plant all taxa in all habitats\u2014is called a reciprocal transplant. I repeated any measurements taken in this garden in the other two, both within a couple hours' drive. A reciprocal transplant is a powerful test for local adaptation. Populations that are adapted to different ecological niches are unlikely to encounter each other in their distinct habitats, and if they do, the nonlocal taxon is likely maladapted and will not persist. Local adaptation, therefore, may contribute to the divergence of closely related lineages. In general, Phlox pilosa subsp. pilosa favors open grassy areas in full sun, while P. amoena grows in the grassy fringes of mixed hardwood forest, and P. pilosa subsp. deamii peppers the understory of similar forest edges. But because the ecological factors that differentiate the preferred habitats of my three Phlox taxa are multidimensional and not entirely obvious to my human senses, I let the wild populations guide me to appropriate sites for the experiment. Settling into my morning work routine, I opened a spreadsheet on my tablet that contained a stack of three-digit codes in a column on the left. Each code corresponded to a unique plant identifier that was stamped into an aluminum tag and fastened in the ground at the base of each plant. In order to test for local adaptation, I designed my experiment to evaluate traits related to fitness, like susceptibility to herbivore damage and total reproductive output. My goal on this visit was to score the presence or absence of herbivore damage and count the number of open flowers on every plant. I labeled two new columns (\"herbivory_2\" and \"flowers_2\") and eased into a cross-legged seat on the edge of my plot. Collecting these data was a comprehensive sensory experience. As I pushed and pulled inflorescences aside to reveal more clusters of bright pink, my fingers reluctantly harvested the sticky secretion that protects the flowering branches of Phlox pilosa subsp. pilosa. Each time I agitated a bunch of flowers, a small flare of sweet fragrance mixed with the sharp scent of spring grasses and forbs soaking in the midmorning sun. The exaggerated buzz of a carpenter bee hummed under the exclamations of chattering songbirds. A jumping spider tickled across my wrist. Sitting quietly, eye-level with the asters (Erigeron philadelphicus), I immersed myself in the dense fabric of interactions that contributed to the deceptively neat figures in my spreadsheet. This common garden, in the full-sun habitat of Phlox pilosa subsp. pilosa, is tucked into the northern tip of a 170,000-acre inland peninsula (the largest in the United States), which spans the border between Kentucky and Tennessee. When the Tennessee Valley Authority completed the two dams that isolated this strip of land, aptly named Land Between the Lakes, the residents were forced to move, leaving their properties to be reclaimed by mixed hardwood forest. The house that complemented this yard and surrounding fields has long since been 4 Arnoldia 77\/4 \u2022 May 2020 demolished, but a patch of feral bearded irises (Iris x germanica) and a single mature post oak (Quercus stellata) hint at where it once stood. These days, the property is mowed annually and burned periodically by the United States Forest Service as part of a scattered network of restored prairie patches, important habitat for the robust deer and turkey populations in this National Recreation Area. My research permit with the local Forest Service office, however, guaranteed that this field would not be burned from spring 2018 through fall 2020, and my garden plot, demarcated with pink marking flags, would not be mowed. The Phlox that I study are the hangers-on of a much more audacious long-term experiment\u2014 the conversion of southeastern prairie into farm and forest. Through the conversion into farmland, suppression of fire, and elimination of grazing bison, humans removed the sources of periodic disturbance that once precluded large trees and favored communities of resilient herbs and grasses. These changes have been compounded by the ebb and flow of fertilizer and pesticide use, an evolving system of hunting regulations, and a rapidly changing climate, creating a volatile experiment with few constant variables. Each species has borne witness to the arc of human impact in its own way. The bald eagle, once suffering, now thrives. The same is true for white-tailed deer and wild turkey. Free-ranging bison have not returned, but Land Between the Lakes supports two small populations of reintroduced bison that graze on fenced-off grasslands, an allusion to the millions that roamed widely until the early 1800s. Dozens of species of prairie-dwelling plant have retreated to small patches of suitable habitat and are threatened or endangered. Today, the closest approximation to the lost prairie disturbance regimes is often the roadside, periodically grazed by a fleet of Department of Transportation mowers. These parallel ribbons are precious refugia for what remain of the remnant prairie species in this part of the world, including the Phlox that I study. Each Phlox in the author's common garden is identified with a numbered aluminum tag, staked at the base. Phlox Common Garden 5 In the Southeast, many grassland species, including the three types of Phlox studied by the author, are confined to roadside strips. Research on the presence and strength of local adaptation may be especially relevant as humans continue to modify the environment. As the southeastern prairies shrank, these three Phlox withdrew into smaller and smaller patches of suitable habitat. If the Phlox were forced into shared fragments, their chances of contacting one another, hybridizing, and melting into one shared gene pool likely increased. Yet, with the exception of a half mile of roadside in western Tennessee, I have never found any of my study taxa living together. After my initial round of spring observations, I would return to these bustling common gardens every few weeks to track flower output as well as the number of fruits each plant produced, the number of seeds in a subset of those fruits, and the aboveground biomass at the end of the growing season. These traits quantify survival, growth, and reproduction, all aspects of fitness that would allow me to test my prediction that these subspecies are locally adapted to distinct ecological conditions. If so, it would help explain how they kept their ecological distance, even as they were concentrated into small pockets of prairie-like habitat. By seven o'clock, the tall grass around me glowed pink. The yellowthroats and gnatcatchers resolved their conversation for the day, and I strained to distinguish the Phlox flowers from one another. I had counted more than six thousand flowers on about two-thirds of the plants in the garden\u2014a tedious but satisfying task\u2014 and would finish the rest before the next day's lunch. I gathered my water bottle and tablet and swished back through the tall grass. Standing next to the rental car, I shook off the tunnel vision of counting flowers and let my eyes wander over the rolling field. My gaze landed again on the large dead tree. A bald eagle leapt from an upper branch, circled the field once, and slipped out of sight behind the canopy. Ben Goulet-Scott is a doctoral candidate in the Department of Organismic and Evolutionary Biology at Harvard University and a fellow of the Arnold Arboretum."},{"has_event_date":0,"type":"arnoldia","title":"Revisiting the Mystery of the Bartram Oak","article_sequence":2,"start_page":6,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25701","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270b36e.jpg","volume":77,"issue_number":4,"year":2020,"series":null,"season":null,"authors":"Crowl, Andrew; Bruno, Ed; Hipp, Andrew L.; Manos, Paul","article_content":"CROWL, A., BRUNO, E., HIPP, A., AND MANOS, P. 2020. REVISITING THE MYSTERY OF THE BARTRAM OAK. ARNOLDIA, 77(4): 6-11 An impressive oak tree grows on the quad of West Chester University, outside of Philadelphia. It is a healthy, open-grown individual measuring approximately 110 feet (33.5 meters) tall and with a trunk diameter of 64 inches (1.6 meters) at breast height. As the oldest tree on campus, it has become an important landmark for students. The tree is also a putative descendant of the first-described Bartram oak (Quercus x heterophylla) and is the largest of its kind in Pennsylvania. As such, the tree was recently recognized as a state champion, but this title remained somewhat uncertain, given the perplexing taxonomic status of the Bartram oak. Ed Bruno, the landscape designer at West Chester University, has been working with the trees on campus for more than thirty years. Bruno was aware of an 1862 observation by the southeastern naturalist Samuel Buckley, indicating that the West Chester oak was perhaps a second-generation descendant of the original Bartram specimen\u2014a seedling of a seedling. The original tree, however, is long gone, which meant that the West Chester oak\u2014now approximately 170 years old\u2014could not be directly compared to it. For Bruno, the identity of the tree became increasingly frustrating. To provide some clarity, Bruno contacted a dozen or so oak taxonomists in 2015, requesting their opinion of the tree's hybrid status and possible ancestry. He shared images of leaves, twigs, buds, bark, and acorns. Most recipients responded with slightly different opinions but agreed the tree was of hybrid origin. The varied answers, however, left the identity in ongoing limbo. Paul Manos, professor of biology at Duke University, agreed with the current identification of the specimen as a possible Bartram oak but suggested DNA testing would be necessary for verification. Testing would also provide an exciting opportunity to finally check hypotheses regarding the putative parents of this famous tree. The results would shed light on a two-hundred-year-old botanical mystery and further the narrative of hybridization as a frequent and important phenomenon in oaks. Revisiting the Mystery of the Bartram Oak Andrew Crowl, Ed Bruno, Andrew L. Hipp, and Paul Manos ANDREW CROWL Facing page: The Bartram oak (Q. x heterophylla, center) displays a range of leaves. Some resemble the willow oak (Q. phellos, top left); others resemble the northern red oak (Q. rubra, top right). These samples are from Duke Gardens. History of the Bartram Oak The original Bartram oak grew near Philadelphia, on the west bank of the Schuylkill River. In the mid-eighteenth century, the tree caught the eye of John Bartram, who was among the first practicing Linnaean-era botanists in the American colonies. Bartram traveled extensively throughout eastern North America, cataloguing and collecting native plants. While the anomalous oak, located within walking distance of Bartram's home, resembled known oak species of the region, it possessed distinct\u2014 though somewhat ambiguous\u2014morphological attributes, such as irregular lobing of the leaves and a range of leaf types from unlobed to lobed. This form of variation is termed heterophylly and likely prevented the specimen from being formally classified for another half century. In 1802, French botanist Fran\u00e7ois Andr\u00e9 Michaux traveled to Philadelphia where he met with John Bartram's son, William Bartram, an accomplished botanist and naturalist in his own right, who was maintaining and growing his father's botanical collection. During this visit, Michaux presumably observed the tree for the first time. When Michaux formally named Quercus heterophylla\u2014coining the common name Bartram oak\u2014in his North American silva, published in 1812, he designated the taxon as a new species rather than a hybrid. Michaux described the morphological ambiguity and suggested that although the Bartram oak resembled the laurel oak (Q. laurifolia), the leaves of that species were never lobed and the closest known population was more than one hundred miles from Philadelphia. The newfound species status bestowed upon the Bartram oak, however, was quickly called into question, in 1814, by Pennsylvanian botanist Frederick Pursh, who had previously served as a horticultural manager at a neighboring estate, known as the Woodlands. \"Of this singular species there is but one individual known, which grows on the plantation of the Messrs. Bartrams near Philadelphia,\" Pursh wrote. \"It probably is only a hybrid plant on that account, and cannot with propriety be considered a genuine species.\" This first suggestion of a hybrid origin was followed by one hundred years of confusion and arguments between botanists as to the validity of this taxon as a distinct species, its hybrid status, and its potential parents. Tragically, in 1842, almost two decades after the death of William Bartram, botanist Thomas Nuttall reported that the original tree had been recently cut down. Thomas Meehan, a preeminent American horticulturist, added to this report in 1853, noting that the tree had been removed because it \"interfered with a view of the Schuylkill [River] from the Woodlands.\" However, acorns of the tree had been collected before the removal and planted on the property and elsewhere around Pennsylvania. In subsequent years, numerous additional examples of this taxon were discovered in New Jersey, Delaware, Maryland, and New York. Though the infamous tree had been lost, this was not the end of the Bartram oak\u2014as a lineage or a botanical mystery. By the mid-1800s, seemingly every notable American botanist, and many from abroad, had examined either an herbarium specimen of a Bartram oak or an actual tree. But the debate continued, and in the words of botanist Arthur Hollick, looking back on this taxonomic foment in 1919, \"The opinions expressed in connection with [the Bartram oak] were as diverse and heterogeneous as the trees were heterophyllous.\" During this period, the Bartram oak was identified by various experts as Quercus ambigua, Q. phellos, Q. imbricaria, Q. laurifolia, Q. hemisphaerica, Q. coccinea, Q. leana, Q. tinctoria (or Q. velutina), Q. aquatica (or Q. nigra), Q. palustris, or some combination of these. A trend did begin to emerge, however, during the latter half of the nineteenth century: the Bartram oak was clearly aligned, in some way, with the willow oak (Quercus phellos). This was based primarily on leaf morphology, with the willow oak exhibiting unlobed and entire leaf margins. Some authors believed the Bartram oak to be a lobed form or variety of the willow oak; others maintained that it was simply an anomalous willow oak specimen; and others (perhaps the majority) argued for a hybrid origin in which the willow oak was a parent. The second parent continued to be debated. Among those that subscribed to the hybrid hypothesis were famed botanists Asa Gray and George Engelmann. Gray, in 1863, expressed Bartram Oak 7 Q. marilandica 3 Q. coccinea 2 Q. inopina 1 Q. marilandica 4 Q. myrtifolia 1 Q. elliottii 1 Q. agrifolia Q. arkansana 2 Q. imbricaria 1 Q. marilandica 2 Q. ilicifolia Q. laevis Q. incana 1 Q. coccinea 3 Q. imbricaria 2 Q. wislizeni 2 Q. buckleyi 2 Q. wislizeni 1 Q. buckleyi 1 Q. marilandica 1 Q. humbodtii Q. canbyi Q. georgiana 1 Q. kelloggii Q. nigra 1 Q. nigra 2 Q. texana 2 Q. arkansana 1 Q. laurifolia 2 Q. hemisphaerica Q inopina 2 Q. ellipsoidalis 1 Q. acerifolia 2 Q. myrtifolia 2 Q. acerifolia 1 Q. texana 3 Q. parvula Q. pagoda 2 Q. georgiana 2 Q. pagoda 1 Q. shumardii 2 Q. palustris 2 Q. coccinea 1 Q. laurifolia 1 Q. palustris 1 Q. ellipsoidalis 2 Q. shumardii 1 Q. elliottii 2 Q. incana 2 Q. mexicana Q. texana 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. velutina 2 Q. velutina 1 Q. rubra 1 Q. rubra 2 Q. falcata 2 Q. falcata 1 Q. x heterophylla Q. phellos 1 Q. phellos 2 Q. phellos 1 Q. phellos 2 Q. falcata 1 Q. falcata 2 Q. x heterophylla Q. phellos 1 Q. phellos 2 Q. rubra 1 Q. rubra 2 Q. x heterophylla H1 H2 H3 Q. phellos 1 Q. phellos 2 Q. velutina 1 Q. velutina 2 Q. x heterophylla * * * Quercus phellos Quercus rubra Quercus x heterophylla * Evolutionary relationships of red oaks (Quercus section Lobatae) are rendered on a phylogenetic tree (left), showing the clear affinity between the Bartram oak and the willow oak (Q. phellos). This complements results from genetic clustering analyses of the Bartram oak and its hypothesized parent species (below). Each colored box represents an individual tree, with colors indicating the genomic composition of that individual. For H1 and H2, the Bartram oak is distinct from the other species tested, while in H3, the genomic content of the Bartram oak demonstrates a clear combination of Q. phellos and the northern red oak (Q. rubra). The map shows the distribution of the Bartram oak and its putative parent species. A black star indicates the location of the West Chester oak. Bartram Oak 9 the opinion that the Bartram oak was a hybrid between the willow oak (Quercus phellos) and black oak (Q. tinctoria, now Q. velutina). Engelmann, on the other hand, disagreed with this and argued, for a time, that the Bartram oak should be considered a distinct species. By 1877, however, he had clearly aligned his thinking with Gray, though he disagreed as to the second parent species: \"While I was long inclined to follow Michaux in considering it as a distinct species \u2026 That it is a hybrid is most probable,\" Engelmann wrote. \"One of its parents is undoubtedly Phellos; for the other we must look among the lobe-leaved Black-oaks of its neighborhood, falcata, rubra or coccinea,\" meaning the southern red oak, northern red oak, and scarlet oak, respectively. At long last, in 1905, nearly one hundred years after Michaux's recognition of the Bartram oak, a group from the New York Botanical Garden attempted to put the debate to rest, once and for all. Arthur Hollick, then the assistant curator of the garden, later reported that seventy-five acorns from a tree on Staten Island had been collected and propagated to test the hybrid hypothesis. The resulting seedlings exhibited considerable variation in leaf morphology, which could be arranged in a series according to the extent of their lobing. On one end of the spectrum were trees exhibiting the deep-lobed leaves of northern red oak (Quercus rubra), while others had narrow leaves with entire margins, similar to those of willow oak (Q. phellos). The remaining individuals were heterophyllous trees, exhibiting various combinations of red and willow oak leaf forms. This was convincing evidence that the two parents for the Bartram oak were Quercus phellos and Q. rubra, and for a long time, this was the only hard evidence regarding the identity of the hybrid. But over one hundred years after this New York Botanical Garden study\u2014and two hundred years after Michaux's account was first published\u2014we reopened the case. This time, however, we had access to DNA sequencing technologies and computational methods, allowing us to peer into the genomes of these trees and directly observe the genetic composition. Modern Investigation In an attempt to shed light on the identity of the West Chester tree\u2014and to provide insights into the background of the original Bartram oak\u2014we broadly sampled North American red oaks, including any species hypothesized to be involved in the hybrid history. We also collected material from the West Chester tree. We then used a genomic sequencing technique (restriction site-associated DNA sequencing or RADseq) to create a genetic dataset for these taxa, resulting in tens of thousands of informative DNA sites for downstream analyses. Based on these data, evolutionary relationships were visualized with a phylogenetic tree. Much like a family tree, a phylogeny is a diagram depicting a pattern of descent and relationships between organisms. It is important to note that the behavior of hybrids in phylogenies is not straightforward and often results in one of two outcomes: the hybrid may be found as a close relative to one of the parent species, or it will be placed in an intermediate position in the tree, falling somewhere between the two parent species. Our phylogenetic analyses confirmed a close relationship of the Bartram oak with willow oak (Quercus phellos). We then carried out additional DNA analyses using a clustering approach that groups individuals based on DNA similarities and differences. This technique can be used to infer the presence of admixed individuals (those whose genomes are a combination of different parent species). Based on previous assertions and our own morphological insights, we tested three plausible hypotheses regarding possible parental lineages: willow oak crosses with southern red oak (Quercus falcata), black oak (Q. velutina), and northern red oak (Q. rubra). Our clustering analyses indicated the genome of this specimen is a mosaic, suggesting a hybrid origin, with northern red oak (Q. rubra) as the probable second parent. This confirmed morphological observations of the 1905 New York Botanical Garden study, as well as our own detailed observations. The West Chester oak shows many fruit characteristics similar to Quercus rubra. Fruit size is larger than would be expected given any of 10 Arnoldia 77\/4 \u2022 May 2020 the other potential parents, measuring up to 1.2 inches (30 millimeters) in length, consistent with the large nuts of Q. rubra. The cup covering of the nut also suggests Q. rubra as a likely parent: while the cup of this taxon covers approximately one-quarter of the nut, Q. velutina and Q. falcata both possess cups that cover up to one-half of the nut. Cup scale arrangement is consistent with Q. phellos and Q. rubra, both of which have smooth and tightly appressed scales. Bud size and bud scales are also consistent with Q. rubra. Leaf pubescence is reminiscent of Q. phellos, which presents hairs early in development but becomes glabrous to sparsely pubescent later in the season. Late-season leaves of the West Chester oak are mostly glabrous, with tufts of hairs in the axils of veins on the underside, much like Q. rubra leaves, which are glabrous throughout development but with similar tufts of hair. Moreover, the bark of the Bartram oak is reminiscent of Q. rubra, with smooth patches on the trunk. This hybrid scenario for the Bartram oak is plausible given the overlapping distributions of willow oak and northern red oak at the edges of their current ranges in eastern North America. As the West Chester tree is likely a secondgeneration offspring of the original Bartram oak, we propose the West Chester tree is the result of backcrossing with willow oak, a common element of the forest in the Philadelphia area. Conclusions and Broader Implications Many questions remain about the Bartram oak due to the inclusion of only a single individual in this study, but the interaction between the two parent species is clear. The parents share only a narrow range of ecological space, yet numerous hybrid individuals have been reported from the northern edge of the willow oak (Quercus phellos) range, distributed in disparate patches. This pattern is likely facilitated by an expanded distribution of willow oak due to land conversion during the last two hundred years, creating increased opportunities for the natural formation of Q. x heterophylla. While we were unable to test whether all Bartram oaks are descendants of a single hybridization event, we believe it to be unlikely. Known Bartram oak specimens are often found as single individuals. In fact, a putative Bartram oak was recently identified by Paul Manos within Duke Gardens, on the campus of Duke University, after years of being noted as an anomaly by garden staff. This single eighty- to ninety-yearold tree occurs, along with both parent species, on the edge of the garden in an area that was historically forested. This suggests the Duke individual is a naturally occurring hybrid rather than an intentional planting. We posit Bartram oaks are the result of multiple independent events that have occurred repeatedly. Future studies with increased sampling will be needed to directly test this hypothesis. Hybridization is certainly a common phenomenon in oaks; however, past concerns of oaks failing to form genetically coherent entities that merit species status have not been substantiated by genetic data. Based on recent DNA studies, we know that oak species have originated by diverging from one another in spite of gene flow. Oak hybrids are known to be fertile, and may eventually participate in forming narrow genetic bridges between species and generating new genetic combinations. This view of species as potentially open systems is based on observations made by generations of botanists. As more organisms across the tree of life are studied, the zoocentric definition of species as reproductively isolated end products of evolution is beginning to fade into history. This new paradigm redirects the question of species status to instead consider the evolutionary potential of naturally occurring Bartram oaks and the role of hybridization, in general, as oaks continue to respond to rapidly changing climates and landscapes. The West Chester oak, in its relative isolation as a prized campus monument, is unlikely to contribute to this evolutionary continuum of gene swapping. But in natural populations, hybridization is no doubt playing a role in shaping the genetic architecture of future generations of trees. For now, and to satisfy those who need to classify and at the same time honor our rich botanical heritage, it seems fitting (and useful) to recognize all first- and later-generation hybrids of Quercus phellos and Q. rubra that show intermediate morphological qualities as Bartram oaks (Q. x heterophylla). And in the meantime, the champion West Chester tree remains a noteworthy destination for anyone with horticultural wanderlust. Bibliography and further readings Buckley, S. B. 1862. Note no. 2: On Quercus heterophylla, Mich. Proceedings of the Academy of Natural Sciences of Philadelphia, 14: 100-101. Conte, J. L. Le, H. Allen, S. B. Buckley, W. M. Gabb, W. Stimpson, and E. Coues. 1861. Note on the Bartram Oak (Quercus heterophylla). In Proceedings of the Academy of Natural Sciences of Philadelphia, 13: 335-390. Philadelphia: Academy of Natural Sciences. Engelmann, G. 1877. The oaks of the United States. Transactions of the Academy of Science of Saint Louis, 3: 539-543. Hipp, A. L., Manos, P. S., Hahn, M., Avishai, M., Bod\u00e9n\u00e8s, C., Valencia-Avalos, S. 2019. Genomic landscape of the global oak phylogeny. New Phytologist. doi:10.1111\/nph.16162 Hollick, A. 1919. The story of the Bartram Oak: How a little exact experimental science solved a problem of long standing. Scientific American, 121(17): 422-432. MacDougal, D. T. 1907. Hybrids among wild plants. The Plant World, 10: 25-27. Martindale, I. 1880. Notes on the Bartram Oak: Quercus heterophylla, Michx. Camden, NJ: S. Chew. Meehan, T. 1853. The American handbook of ornamental trees. Philadelphia: Lippincott, Grambo, and Company. Meehan, T. 1902. Contributions to the life-history of plants, No. XVI. Proceedings of the Academy of Natural Sciences of Philadelphia, 54: 33-36. Michaux, F. A. 1812. Histoire des arbres forestiers de l'Am\u00e9rique septentrionale, consid\u00e9r\u00e9s principalement sous les rapports de leur usages dans les arts et de leur introduction dans le commerce. Paris: L. Haussmann. Nuttall, T. 1842. The North American sylva, or, a description of the forest trees of the United States, Canada, and Nova Scotia, not described in the work of F. Andrew Michaux. Philadelphia: J. Dobson. Pursh, F. 1814. Flora Americae Septentrionalis; or a systematic arrangement and description of the plants of North America (Vol. 2). London: White, Cochrane, and Company. Trelease, W. 1917. Naming American hybrid oaks. Proceedings of the American Philosophical Society, 56: 44-52. Andrew Crowl is a postdoctoral associate at Duke University. Ed Bruno recently retired as landscape designer at West Chester University, after a thirty-year career with the school. Andrew Hipp is senior scientist in plant systematics and herbarium curator at the Morton Arboretum. Paul Manos is a professor in the Department of Biology at Duke University. The Bartram oak on the campus of West Chester University retains its claim as a state champion. ED BRUNO"},{"has_event_date":0,"type":"arnoldia","title":"Collector on a Grand Scale: The Horticultural Visions of Henry Francis du Pont","article_sequence":3,"start_page":12,"end_page":23,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25703","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270b76b.jpg","volume":77,"issue_number":4,"year":2020,"series":null,"season":null,"authors":"Wilkie, Carter","article_content":"In 1924, the Arnold Arboretum's first director, Charles Sprague Sargent, named a new hybrid buckeye, Aesculus x dupontii, in the Journal of the Arnold Arboretum and praised the tree's namesake, the du Ponts, for making the vicinity of Wilmington, Delaware, \"one of the chief centers of horticulture in the United States.\" The family's fortune had exploded from the manufacture of gunpowder in the nineteenth century and was enriched further by chemicals in the twentieth. The resources the du Ponts dedicated to their landscapes made Delaware's Brandywine Valley a must-see destination for horticulturists who travel there to visit and study at the estates that are open to the public today. Longwood Gardens, just over the Delaware border in Kennett Square, Pennsylvania, is the most visited, with its Italianate and French neoclassical fountains, lightshows, and fireworks that elicit audible \"oohs\" and \"ahhs\" from large crowds at all seasons of the year. Its creator, Pierre S. du Pont, was inspired by the spectacle and sense of wonder he experienced, at age six, when he attended the Centennial Exhibition in Philadelphia in 1876. His grand conservatories of hothouse plants continue to wow his posthumous guests, especially when decorated for Christmas, as does a garden amphitheater that hosts evening concerts and Shakespeare plays performed in a veritable Forest of Arden. Closer to Wilmington, his cousin Alfred I. du Pont spent a large fortune to build Nemours, with gardens modeled after Versailles. And Mt. Cuba, the estate of Lammot du Pont Copeland and his wife, Pamela, would become a display garden and research center for studying the native flora of the Piedmont. Of the estates that earned northern Delaware the sobriquet \"chateau country,\" Henry Francis du Pont's Winterthur Museum and Gardens is the most naturalistic. Home to nearly a thousand acres of rolling meadows, forests, and one of the finest woodland gardens in the world, Winterthur's connections to the Arnold Arboretum are deep. To walk the curving pathways through its woods and fields is to see a landscape shaped by what H. F. du Pont learned in Boston and through collaboration with the Arboretum's collectors and propagators over decades. In an affectionate yet frank book about Winterthur (pronounced \"winter tour,\" meaning \"winter's door\"), H. F. du Pont's daughter Ruth Lord claimed that her father found his life's calling at the Arboretum. As a student in his junior year at Harvard, in 1901, du Pont applied for admission into classes at Harvard's Bussey Institution, one of the first formal university programs to teach horticulture in America. Its mission, according to the Bulletin of the Bussey Institution, was to educate \"young men who intend to become practical farmers, gardeners, florists, or landscape gardeners,\" as well as \"men who will naturally be called upon to manage large estates.\" Young du Pont was destined to become all of those things. But by October, the fall semester had already begun. He was late and had an unimpressive academic transcript. The coursework was rigorous, taught by scientists with little patience for dilettantes. Still, he was admitted with the expectation that he could catch up. He wrote to his mother of his \"sudden resolution \u2026 my great desire to really know something about flowers \u2026 In fact flowers etc. are the only real interests I have.\" He added, \"I do not think I am impulsive I hope not at least. I merely think it is the smouldering [sic] of latent thought which has burst into flame.\" In his first course, Horticulture I, Collector on a Grand Scale: The Horticultural Visions of Henry Francis du Pont Carter Wilkie Facing page: The sweeping and naturalistic landscape of Winterthur Museum and Gardens was shaped by the horticultural vision of Henry Francis du Pont. ALL PHOTOS BY THE AUTHOR UNLESS NOTED 14 Arnoldia 77\/4 \u2022 May 2020 taught by Benjamin Marston Watson, who led Harvard's horticultural instruction program for almost forty years, du Pont received a D-. The student would turn out to be a late bloomer. From that unpromising beginning, H. F. du Pont went on to become one of the most accomplished horticulturists of the twentieth century, a man the Garden Club of America in 1956 designated as perhaps \"the best gardener this country has ever produced\" up until that time. He also served as an important benefactor of the Arboretum and would consult with its staff over the next seven decades. Scion of a Distinguished Family Tree H. F. du Pont (or \"Harry,\" as his family called him) was born in 1880, \"with a silver trowel in his hand.\" He was the son of Henry Algernon du Pont, the richest man in Delaware; grandson of Henry du Pont, the longest-serving chief executive of the E. I. du Pont de Nemours Company; and great-grandson of the company's founder, \u00c9leuth\u00e8re Ir\u00e9n\u00e9e du Pont, who arrived in America with three generations of du Ponts in 1800. E. I. du Pont settled his family and established his powder works on the bank of Delaware's Brandywine River. There, at what he named Eleutherian Mills, he laid out a French parterre, with fruit orchards and potager to feed his family. He imported trees from Europe and instilled in his children and grandchildren a love of horticulture and an interest in agriculture and animal husbandry. E. I. du Pont purchased the first acres of what would become Winterthur with gunpowder profits from the War of 1812. H. F. du Pont's father inherited the property in 1889. By then, Winterthur had sprawled to 1,135 acres. As children, H. F. du Pont and his older sister, Louise, had the run of the outdoors, with farm animals for companions: goats, sheep, poultry, and forty draft horses. In her late seventies, Louise recalled to Winterthur curator John Sweeney how her father drilled into them his interest in botany, and the process of learning through careful observation: \"Father would take Du Pont studied horticulture at Harvard's Bussey Institution, which was located on South Street, adjacent to the Arnold Arboretum. ARNOLD ARBORETUM ARCHIVES Henry Francis du Pont and Winterthur 15 Harry and me by the hand and walk through the gardens with us, and if we couldn't identify the flowers and plants by their botanical names, we were sent to bed without our suppers.\" If their father, first in his class at West Point in 1861, was pompous and rigid, their mother, Pauline Foster, was warm and tender. Having lost five of seven children in infancy, she kept her son close and passed on to him her love of flowers that she shared with her mother-in-law, Louisa Gerhard du Pont, and other du Pont relatives. Pauline was the daughter of a gentleman farmer in New York, and she impressed upon her son that Winterthur was not a showplace but a country place, a retreat for repose. H. F. du Pont was shy as child and awkward around peers in his youth (he spoke only French when he first entered school), and he would credit his mother with his lifelong desire to reinforce the feeling of \"great calm and peace\" that Winterthur provided in his anxious childhood. At age thirteen, H. F. du Pont was sent off to boarding school at Groton, Massachusetts. From letters to his parents, he hated being away and consoled himself with visual memories of home. He wrote of his joy at recognizing Winterthur's May-blooming Brandywine bluebells (more commonly known as Virginia bluebells, Mertensia virginica) in Gray's Manual of Botany. When he begged for permission to work in the nurseries of a garden center near the school, du Pont's parents fretted over their son of the manor getting his hands dirty and rubbing elbows with workingmen in Groton. But he had already performed chores for Winterthur's gardeners, who decades later would remark that du Pont could work as hard physically as any paid laborer. Student at the Bussey Institution After entering Harvard in 1899, du Pont reconnected with a childhood acquaintance, Marian Coffin, one of two women enrolled in the new landscape architecture program at the Massachusetts Institute of Technology. (Harvard did not admit women at the time.) Coffin's mother and du Pont's mother were close friends. It was Coffin who had urged du Pont to take courses at the Bussey Institution. Together, they studied in the Arboretum under John George Jack and toured Holm Lea, Sargent's 150-acre estate in Brookline, on the north side of Jamaica Pond. There, du Pont expected to find mature trees of enormous size but wrote home with disappointment that he saw only two, although he noted that \"the Magnolias around the pond were in full bloom and magnificent.\" Coffin found in Sargent a mentor who had already taken under his wing the early female pioneer in landscape architecture, Beatrix Jones (Farrand). Coffin's program at MIT, under the direction of Sargent's son-in-law Guy Lowell, emphasized geometric gardens in the neoclassical tradition. Homeschooled before college, Coffin found the heavy math requirement daunting. She credited Sargent with encouraging her to persevere, effectively saving her career at a critical moment of self-doubt. The death of du Pont's mother in his junior year made him return home and spend his senior year helping his father run the household and its staff. After graduating, Coffin and du Pont would tour the great gardens of Europe together, with her mother as chaperone. In an era when few firms would hire a woman landscape designer or have one supervise all-male crews, Coffin struck out on her own. Du Pont, meanwhile, would become a valuable client and steer business her way. He put off planned studies in New York's Hudson Valley, at the School of Practical Agriculture and Horticulture in Briarcliff Manor, and began to apply at Winterthur the knowledge he had acquired at the Bussey Institution, experimenting with plants, observing how they performed, and carrying with him a notebook everywhere he went. He started a trial of fifty-four different daffodils and planted the ones that performed best (Narcissus horsfieldii, N. albicans, and the cultivars 'Golden Spur', 'Grandee,'' and 'Emperor') along the banks of a stream and on hillsides, in large drifts and massed colonies, never mixing them. A Bussey course on hardy herbaceous plant materials had introduced him to the ideas of William Robinson, the evangelist for naturalistic gardens, whose book The Wild Garden, published in 1870, had revolutionized landscape design in Britain. The Irish-born Robinson was an irreverent crusader against Victorian garden contrivances, from the bedding out of 16 Arnoldia 77\/4 \u2022 May 2020 tender, tropical annuals in temperate climates to the idolatry of faux Italianate ruins. Instead, Robinson advocated for the use of winter-hardy plants and natural-looking gardens \"devoid of any trace of man.\" Valencia Libby, who dug deeply into Winterthur's ties to the Arboretum, unearthed a paper that du Pont wrote at Harvard (about an aunt's estate, Virieux, which bordered Winterthur) that reveals Robinson's strong influence on an impressionable student. Robinson had also influenced Sargent and the Arboretum's original landscape architect, Frederick Law Olmsted. Laconic rather than loquacious, du Pont never articulated his design principles in one comprehensive place for easy retrieval. Scholars have pieced them together from snippets he offered here and there and from the visual evidence he left behind. Above all, he strove to achieve the appearance of nature working effortlessly, with COURTESY OF THE WINTERTHUR LIBRARY: WINTERTHUR ARCHIVES H. F. du Pont relaxes in the Winterthur landscape, in 1904, with Marion Rawson and cousin Elaine Irving. Henry Francis du Pont and Winterthur 17 the garden fitting into the landscape \"as if it has always been there,\" he would say. He told visitors this design effect took a great deal of effort and was \"very hard to do.\" Estate Planner, Arboretum Benefactor In 1909, when du Pont's father gave him control of the estate's grounds and greenhouses, the young horticulturist began acquiring plants with the zeal of an obsessive-compulsive collector on an unlimited budget, planting twenty-nine thousand bulbs that year and thirty-nine thousand the next. He carpeted the ground beneath tulip poplars (Liriodendron tulipifera) with snowdrops (Galanthus), winter aconite (Eranthis hyemalis), glory-of-the-snow (Chionodoxa luciliae), squill (Scilla), snowflake (Leucojum vernum), and crocus (Crocus tomasinianus). In the decades to follow, he would source bulbs, herbaceous perennials, and woody plants from the top breeders and more than fifty nurseries, chief among them the Arnold Arboretum. Over the years, Sargent evolved from du Pont's professor and advisor into a peer and beneficiary. The du Ponts appear in Sargent's annual reports on the Arboretum to Harvard's treasurer beginning in 1915, when du Pont's father, Henry A. du Pont, made a donation to fund annual operating expenses, the equivalent of almost $2,500 today. Sargent, sensing an opportunity to cultivate new patrons to sustain the institution, pursued a personal relationship. He made personal visits to Winterthur, signing its guest book nine times between 1918 and 1923, usually during April, when spring in Wilmington is in full bloom while Boston is still dreary. Sargent already knew Wilmington as the home of the wealthy botanist William Canby, who collected forty-five thousand botanical specimens in his lifetime and had accompanied Sargent and John Muir on a tour of the Appalachians. Sargent grew close to H. A. du Pont, hosting him for personal tours of the Arboretum and the Hunnewell Estate, in Wellesley, where Horatio Hollis Hunnewell had popularized the cultivation of rhododendrons, especially the red torch azalea (Rhododendron kaempferi) that Sargent had brought over from Japan. Both Sargent and H. A. du Pont were veterans of the Civil War and patrician practitioners of noblesse oblige. In the twilight of his life, H. A. du Pont called Sargent his favorite friend. In 1916, H. F. du Pont wed the more outgoing Ruth Wales, who had grown up in New York near his former schoolmate at Groton and Harvard, Franklin D. Roosevelt. The next year, Sargent wrote a letter inviting du Pont, the one-time Bussey student, forty years his junior, to serve on the Arboretum's governing committee, formally called the Harvard Board of Overseers' Committee to Visit the Arnold Arboretum. \"The committee appointed by the Overseers has been of very great service to me now for many years in aiding [and] \u2026 in raising enough money \u2026 to keep the establishment going, the income from the endowment being inadequate for that purpose,\" Sargent wrote. \"While the Committee has been of great service to the Arboretum in this way I have never gotten any horticultural or other advice from its members, and when I suggested to the overseers to appoint you as a member of the Committee it was with the idea that you should be able to help me horticulturally for in this direction I am left entirely without advice or assistance.\" Du Pont would serve the Arboretum in that capacity for fifty years, until 1968, the year before he died. Over the years, du Pont would rely upon the Arboretum's experts for plant identification and sourcing, consulting its long-time propagator Jackson Dawson, Dawson's successor William Judd, and later director Karl Sax. With the Arboretum dependent on donors for fundraising, Sargent was more solicitous in correspondence than his staff, replying to one of du Pont's inquiries about the fragrant, white-blooming mock orange (Philadelphus) by writing, \"If there is any particular kind you want, we shall be glad to have a plant propagated for you.\" Du Pont would return to the Arboretum again and again, always with a notebook in hand, on frequent visits to see his sister who lived with her husband, Frank Crowninshield, in Boston and Marblehead. After a visit in 1923, du Pont wrote Sargent: \"The only trouble in going to the Arboretum is that I come back fired to possess all kinds of plants which, as you 18 Arnoldia 77\/4 \u2022 May 2020 know, are unprocurable elsewhere. After going through numberless catalogues I am absolutely stumped by the enclosed list, and I am wondering if little by little you could procure cuttings of these various shrubs, as I should so much like to have them.\" During du Pont's first year on the Overseers' Committee, Henry Hunnewell, son of Horatio Hollis Hunnewell, initiated a capital campaign to grow the Arboretum's endowment. Du Pont and his father made gifts equivalent to six figures in today's dollars. Then, in 1918 and 1919, they each provided the Arboretum with its largest annual financial gifts from individuals. The Great War in Europe had been lucrative for the family's munitions business, and their wealth had multiplied thanks to three of du Pont's second cousins (Pierre S., Alfred I., and T. Coleman du Pont) who had taken control of the DuPont company and engineered its expansion into chemicals and a large stake in General Motors. H. F. du Pont would plow his share into developing Winterthur, collecting antiques, and creating a summer place for his wife in Southampton, New York. Woodland Gardener Before the Great War, du Pont and his father visited a conifer forest that was cultivated at the Dropmore estate, in Buckinghamshire, England. The estate dated to the eighteenth century, and some of its coniferous trees had been propagated by seed brought to England by early collectors. The sight of the rare specimens inspired du Pont's father to install a conifer collection at Winterthur. It grew to contain more than fifty different conifers recommended by Sargent and sourced by the Arboretum and thirteen commercial nurseries. Japanese umbrella pine (Sciadopitys verticillata), Japanese cedar (Cryptomeria japonica), and Atlas cedar (Cedrus atlantica) were among the selections. When laden with freshly fallen snow, Henry Algernon du Pont's dark-green Pinetum becomes Winterthur's own winter's door. While H. A. du Pont installed the Pinetum, his son worked on what would become his crowning outdoor achievement: Azalea Woods. Beginning in early spring and continuing into early summer, eight acres of second-growth tulip poplar, white oaks (Quercus alba), American beech (Fagus grandifolia), and hickories (Carya ovata) are brightened, at eye level, with hues of white, pearl, blush, pink, and red. The design was not conceived at once in any grand plan but grew organically, like seeds sprouting in niches of opportunity opened by the demise of Winterthur's American chestnut (Castanea dentata), killed by blight. Among the azaleas H. F. du Pont used were seventeen Kurume hybrid azaleas he purchased during a visit to Cottage Garden Company, on Long Island. The nurseryman Robert Brown had obtained them from Yokohoma Nursery Company, in Japan, which had won a gold medal for showing them at the San Francisco Exposition of 1915. (The Yokohoma nursery had also grown the bonsai collection that Larz Anderson acquired in 1913 and which his widow donated to the Arboretum in 1937.) At Winterthur, du Pont was delighted when the new azaleas bloomed a subtle shade of pink. From these original accessions, Winterthur propagated more until their progeny spread for acres. In 1920, when Sargent wrote to du Pont with excitement about a new azalea introduction that held great promise for the nursery trade (a group of Kurume azaleas that Ernest Henry Wilson had selected from Akashi Kojiro\u00af, a nurseryman in Kurume, Japan), du Pont modestly avoided telling Sargent that he had been working with Kurume hybrids for three years already. From the Arboretum came Hunnewell's Rhododendron kaempferi, and Sargent recommended royal azalea (R. schlippenbachii), praising it as \"the loveliest of the hardy Asiatic Azaleas.\" In the 1930s, du Pont added broadleaved rhododendron hybrids from Charles Dexter of Sandwich, Massachusetts. Today, Winterthur's plant database catalogues thousands of azaleas on the property, representing 252 species and varieties. The collection reaches peak bloom in Wilmington around the same time as Lilac Sunday at the Arboretum (Mother's Day, the second Sunday in May). In bloom, the shrub layer stands out against the tall trunks and their drab bark the way Boston's sleek John Hancock Tower, designed by I. M. Pei's partner Henry Cobb, plays off of the hefty brown masonry of Henry Hobson Richardson's Trinity Church. Without Trinity Church beside Henry Francis du Pont and Winterthur 19 Azalea Woods was one of the first landscapes that du Pont designed at Winterthur. It has proved an enduring masterpiece. it, the modern glass tower could be an unmemorable building in almost any suburban office park. Likewise, without Winterthur's trees rising out of the shrub layer like giant columns, Azalea Woods would be just azaleas, an overscaled, formless mass of color, lacking apparent depth. At the herbaceous layer, du Pont again followed Robinson's ideas and planted great white trillium (Trillium grandiflorum), blue anemone (Anemone apennina), bluebells, bloodroot (Sanguinaria canadensis), lily of the valley (Convallaria majalis), smaller narcissi, and ferns to naturalize in colonies. While novice gardeners can be reluctant to uproot what they install, du Pont was a ruthless editor of his own work. He was a perfectionist about form and color and personally supervised the installation of trees and shrubs. In oral histories, his gardeners recalled how he would have them move a shrub mere inches to site it perfectly. He would have them plant and replant some shrubs five or six times until everything was right. Color dictated what went where. \"For me, color is the thing that really counts more than any other,\" he told an interviewer at age eighty-two. Viewed through a wide-angle lens, du Pont used color to emphasize the movement of bloom sequence, which rolls across the gardens at Winterthur like slow-moving, undulating waves. By grouping flowering shrubs, he strove for harmony of related hues, or complementary 20 Arnoldia 77\/4 \u2022 May 2020 colors at opposite ends of the color wheel. A signature color combination was mauve against chartreuse, which he produced by coupling two early blooming woody plants: the greenishyellow blooms of winterhazel (Corylopsis glabrescens) with the Korean rosebay rhododendron (Rhododendron mucronulatum). He also brought outdoor colors inside the mansion, decorating rooms with fabrics and cut flowers to reflect what was visible through each window. Guests who arrived at Winterthur for the first time were bowled over by the volume of cut flowers in the public rooms. In the dining rooms, he matched table linens with the flowers and kept more than fifty patterns of china (not place settings but entire sets of china) to do the same with dinnerware. For decades, he kept meticulous notes on every table setting so that returning guests could be served on china they had not seen on previous visits. Collector on a Grand Scale The estate du Pont inherited in 1927, at age forty-six, spread to 2,600 acres. It contained ninety houses for the 250 or so employees working at the estate's mansion, gardens, and farms. The self-supporting community had its own railroad station, post office, a vast complex of twenty greenhouses and potting sheds, cold frames covering an acre, huge livestock barns, a sawmill, tannery, and dairy. Descended from wealthy gentlemen farmers on both \"I like to see the shape and size of big shrubs,\" du Pont would write. Here azalea masses drift beneath conifers at Winterthur Museum and Gardens. Henry Francis du Pont and Winterthur 21 sides of his family, du Pont had been managing all farm operations for thirteen years. On legal documents that asked for his occupation, he sometimes wrote \"farmer.\" He even achieved fame for breeding a champion herd of milking Holstein Friesians, which won top awards from the dairy industry. He raised sheep and poultry, and his daughter remembered how he also loved his pigs. Having full control to shape Winterthur to his liking, one of the first things du Pont altered was his father's Pinetum. To the son, it felt like a collection of specimens arranged artificially, so he naturalized it with quince (Chaenomeles) planted along its broad path, creating his Quince Walk. He gave shrubs room to grow to their natural form. In his single-paragraph foreword for Hal Bruce's 1968 book, Winterthur in Bloom, du Pont echoed Robinson and Sargent, writing, \"I like to see the shape and size of big shrubs; even though they are always part of a group, one has to know when planting just how big and tall the shrubs are going to be.\" Against the dark greens of the conifers he also planted Winterthur's boldest flame azaleas (Rhododendron calendulaceum), which bloom in tangerine, apricot, salmon, and lemon yellow. Later, he added a dawn redwood (Metasequoia glyptostroboides) from the Arboretum. Du Pont then enlarged the big house, which grew to 175 rooms, to accommodate his expanding collection of American antiques. According to his daughter, du Pont's interest in early Americana was sparked by a visit to the Webb estate, in Shelburne, Vermont, in 1923, when he spotted pink Staffordshire china arrayed on a brown pine dresser. Those very pieces are now displayed among the ninety thousand objects of decorative art in Winterthur's collection. A visit that same year to the Gloucester, Massachusetts, home of Henry Davis Sleeper (now owned by Historic New England) inspired du Pont to install period rooms lifted from colonial era houses, as Sleeper had done, and as Ben Perley Poore had done in a haphazard way before at his estate, Indian Hill, in nearby Newburyport. Du Pont's genius as a designer of naturalistic landscapes shows in the way he fit his mansion into the existing topography. He left the north elevation of the existing house at four stories, but on the opposite side, he tucked nine new stories into a steeply sloping hillside that absorbed the height and volume. He also sited the building and new entrances carefully within an envelope of mature oaks, beech, and poplars. The height of the trees, at 150 feet, made the mansion appear less large. From Robinson and Olmsted, he had learned to subordinate built structures to their natural surroundings. As he expanded the house, du Pont hired his friend Marian Coffin to makeover the gardens along its southern shaded slope, which cradles a swimming pool and twin pool houses. The gardens Coffin designed were the most formal at Winterthur, arranged on straight axes punctuated at the ends by semicircles. Her talents complemented his. She architected the skeleton, and he fleshed it out. He outfitted the design with plants, softening her stonework and straight lines with shrubs and understory trees that he allowed to grow naturally, out over the edges. As collaborators, Coffin and du Pont would bounce ideas off one another throughout their lives. Du Pont wrote her playfully during their work on the project, \"I am enclosing a copy of a letter from Mr. E. H. Wilson of the Arnold Arboretum in regard to the Picea asperata notabilis. This is the tree which you wished so ruthlessly to destroy.\" Native to Sichuan, China, dragon spruce (Picea asperata var. notabilis) was described by Wilson and Arboretum taxonomist Alfred Rehder, in 1916, and is considered endangered today. Du Pont weaved Coffin's formal gardens into Winterthur's naturalistic grounds by dissolving boundaries within the landscape. Like Olmsted's design of the Arboretum, there are no obvious seams between garden areas at Winterthur, only gentle transitions. The edge of Azalea Woods dissolves into the meadow beyond it, with shrubs extending out from under trees like an irregular line of troops beginning their advance on an open field. Following Robinson's dictate, Winterthur allows the lower limbs of trees to grow into the ground naturally, eschewing cuts in turf for neatly delineated beds where field and forest meet. Whereas du Pont's cousin Pierre, at Longwood Gardens, had used princess trees (Paulownia tomentosa) to line a formal all\u00e9e to the entrance of his monumental conser22 Arnoldia 77\/4 \u2022 May 2020 vatories, H. F. du Pont inserted Paulownia into his woodland edge as a transition element, its lavender blossoms catching the eye and leading visitors to the next sequence of seasonal bloom. A large mound of saucer magnolia (Magnolia x soulangeana), planted by du Pont's father in 1880, the year du Pont was born, carries the spring bloom into fields of grass, as do two large Sargent cherries (Prunus sargentii) beyond them, gifts to Winterthur from Sargent in 1918. Nearby, du Pont collaborated with Coffin again to create an April-blooming garden of fragrant ornamental trees and shrubs, many of which came from the Arboretum through its Cooperative Nurserymen program. Planner for Posterity In 1930, as du Pont's sister, Louise, planned for the future of Eleutherian Mills, the family's restored ancestral home, H. F. du Pont established a nonprofit entity charged with maintaining Winterthur in perpetuity as \"a museum and arboretum for the education and enjoyment of the public.\" The museum opened to the public in 1951, when du Pont relinquished responsibility for it to professional staff. He continued to oversee the gardens and farmland, calling himself head gardener. As he grew older, du Pont became more impish in the garden. He relaxed his highbrow standards of what constituted good taste. Before mod fashion in the 1960s made pink and orange a popular color combination, he inserted salmon blooming azaleas as accents into his Azalea Woods and placed bold, red-blooming azaleas next to lavender ones to \"chic it up,\" in his words. Coffin praised what she called his \"near discords\" of color. Tossing aside rules he learned about cool, pastel subtlety from the teachings of Gertrude Jekyll, he installed a In 1929, du Pont commissioned his friend Marian Coffin to redesign the south-facing slope below his mansion. Du Pont softened the formal lines with masses of shrubs. Henry Francis du Pont and Winterthur 23 carnival of hot colors in his summer Quarry Garden using primroses (Primula). Gordon Tyrell, who worked closely with du Pont in the garden, confided in a colleague, \"He was mixing colors. I know he did it intentionally, but they were beginning to yell. There were lavenders and mauves and reds. It wasn't offensive, but I think it was a little joke of his really. And I said, 'You can't do this.' And he said, 'I'm doing it.' And he did it.\" Although du Pont loosened Jekyll's tether on color, the aging gardener remained devoted to William Robinson's naturalistic aesthetic into the ninth and final decade of his life. When he hired architects to design a pavilion that became Winterthur's visitor center, he told them, \"Make it look like it isn't there.\" Tucked within his woodland, the modernist building is the color of bark, and its glass exterior walls reflect the foliage around it, camouflaging its mass. The approach road to the visitor center follows swales around hills, through open meadows. He had the road sunken below sightlines so as not to mar the views. Coffin liked to tell prospective clients that great gardens require three things: money, manure, and maintenance. Winterthur had all three in abundance. After du Pont's death in 1969, his endowment supplied the money, but the manure had to come from elsewhere. His will stipulated that his livestock operations be liquidated upon his death, to focus resources on the museum. Because du Pont oversaw and financed the gardens until he died, it took the institution two decades to formalize a Garden Department to preserve his landscape design intent. By then, his naturalistic garden was overgrown. Three years were spent assessing what was there and culling what shouldn't be, including forty truckloads of branches pruned from Azalea Woods. In the Pinetum, a mature Atlas cedar (Cedrus atlantica 'Glauca') was pruned at its base to reopen the circular seating area and sightlines around it. The spot reminds a visitor of the vantage point atop the Arboretum's Bussey Hill Overlook, where puddingstone boulders in the ground encircle the base of a large eastern white pine (Pinus strobus) and Japanese white pines (Pinus parviflora). At Winterthur, Sargent's role in shaping the Pinetum is memorialized on a plaque. Decades before, when du Pont planted bulbs by the tens of thousands, he wrote to Coffin that no mere mortal could do what he wanted done at Winterthur. And he was right. What makes Winterthur unique in America is its scale\u2014the product of vast wealth, space, and time. Few landscapes in the new world are the work of generations of a single family, let alone one man's lifetime of eighty-nine years. Today, Winterthur fulfills Henry Francis du Pont's wish \"that the museum will be a continuing source of inspiration and education for all time, and that the gardens and grounds will of themselves be a country place museum where visitors may enjoy as I have, not only the flowers, trees and shrubs, but also the sunlit meadows, shady wood paths, and the peace and great calm of a country place which has been loved and taken care of for three generations.\" References Bruce, H. 1968. Winterthur in Bloom: Winter, Spring, Summer, Autumn. New York: Chanticleer. Cantor, J. 1997. Winterthur. New York: Harry N. Abrams. Fleming, N. 1995. Money, Manure & Maintenance. Weston: Country Place Books. Libby, V. 1984. Henry Francis du Pont and the Early Development of Winterthur Gardens, 1880- 1927. Master's Thesis for the Longwood Graduate Program in Public Horticulture, The University of Delaware. Lord, R. 1999. Henry F. du Pont and Winterthur: A Daughter's Portrait. New Haven: Yale University Press. Mangani, D., et al. 1995. The Winterthur Garden: Henry F. du Pont's Romance with the Land. New York: Harry N. Abrams. Robinson, W. 1870. The Wild Garden: Or, Our Groves and Shrubberies Made Beautiful By The Naturalization Of Hardy Exotic Plants. London: John Murray. (Also, an expanded edition with new chapters and photography by Rick Darke. 2009. The Wild Garden: Expanded Edition. London: Timber Press.) Carter Wilkie grew up near Winterthur, where his mother took him on frequent visits and would quiz him to identify plants she had pointed out on previous walks. For thirty years, he has resided within a short walk of the Arboretum, where he can be spotted reading tags on trees. This article is based on a talk he gave to the local garden club in Roslindale in March 2018."},{"has_event_date":0,"type":"arnoldia","title":"Eternal Forests: The Veneration of Old Trees in Japan","article_sequence":4,"start_page":24,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25704","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270bb6e.jpg","volume":77,"issue_number":4,"year":2020,"series":null,"season":null,"authors":"Moore, Glenn; Atherton, Cassandra","article_content":"The accounts of foreign visitors who began arriving in Japan after the Meiji Revolution, in 1868, show that the newcomers were intrigued by the Japanese relationship with trees. Sacred trees were noted as important features around shrines. Old trees were marveled over, especially given the affectionate care the trees received, as were the miniature bonsai that could be hundreds of years old and require daily tending. But the visitors focused especially on the cherry trees (Prunus serrulata) and their brief but spectacular show of blossoms. Writer Lafcadio Hearn was no exception, and he recorded fables like \"The Cherry Tree of the Sixteenth Day\" in his classic book Kwaidan, published in 1904.1 Tourists today have the same focus, resulting in the peak season for tourism to Japan happening in early April, when the cherry trees are in bloom. Massed blossoms are the perfect photo opportunity\u2014they accounted for a massive twenty-nine million Instagram posts in 2018\u2014and the cherry-blossom-viewing ritual, known as hanami, is an attraction in itself. The ritual involves siting on blue tarpaulins beneath the trees and drinking sake while the fragile blossoms fall. This is promoted to tourists as indicative of a society whose people accept \"the fleeting nature of life.\"2 Although the symbolism of the cherry blossom certainly seems to fit with a nation Eternal Forests: The Veneration of Old Trees in Japan Glenn Moore and Cassandra Atherton Old Trees in Japan 25 Facing page: Branches of old Japanese red pines (Pinus densiflora) are supported with props in a Kyoto park. PHOTO BY STEPHEN SCHNEIDER that has endured earthquakes, tsunamis, and typhoons, hanami is just a small part of how trees fit into traditional Japanese culture. This relationship is rooted in Japanese history, folklore, and religion. Trees and nature are central to Shinto, a religion that originated in Japan, which holds that spirits inhabit trees that reach one hundred years of age. These tree spirits are known as kodama, and according to Japanese folklore, the kodama give the tree a personality. 3 Accordingly, in premodern Japan, old trees were regarded with awe and a degree of caution. In fact, they were marked with a sacred rope called a shimenawa, warning that if anyone chopped down the tree, they would have to deal with an angry spirit. Although it seems unlikely that these folkloric beliefs could survive in modern, urbanized Japan, they proved surprisingly adaptable. The idea of tree spirits was kept alive in storybooks and anime (most notably the Hayao Miyazaki film Princess Mononoke), and belief in the presence of kodama in old trees continues. So, while it might be more accurate to say that many of today's Tokyoites are likely to think in terms of old trees having admirable qualities rather than personalities, they nevertheless respect the trees for their age and resilience, and can be seen paying homage to them at shrines and in parks, or simply spending time in their presence as an antidote to the stress of modern life. Eternal Forest in Tokyo Tokyo is a modern city, and to a visitor, its residents seem totally immersed in their busy, modern lives. Salarymen dressed in suits and ties rush from train stations to their offices; young people sit in coffee shops engrossed in their phones; and people shop in gleaming department stores. It is easy to lose sight of the fact that Japanese traditions and myths persist behind what writer and cultural historian Boy\u00e9 De Mente refers to as a \"Western fa\u00e7ade.\"4 While not everyone today believes in the ancient myths in a literal sense, Shinto master Motohisa Yamakage has described how myths and related Shinto beliefs are still woven into the fabric of everyday Japanese life.5 For example, construction crews typically wait for a Shinto priest to purify a new worksite; major league baseball teams like the Hiroshima Carp receive a Shinto blessing before spring training; and almost everyone visits one of Japan's eighty thousand Shinto shrines on ceremonial occasions. It is during these shrine visits that the role played by trees in connecting people with the nation's mythic past becomes apparent. Meiji Jingu\u00af , a shrine in Tokyo, is not old by Japanese standards. It was completed in 1920 to honor Emperor Meiji and Empress Sho\u00afken. All Shinto shrines are surrounded by trees, which are thought to provide a conduit to the gods. While these shrine forests, or chinju no mori, are ideally \"old primeval forests,\" providing a living link to \"the ancient age of myths,\"6 everything at Meiji Jingu\u00af had to be planted from scratch, which required a staggering one hundred thousand trees. The long-term goal was to create an \"eternal forest\" dominated by long-lived trees like zelkova (Zelkova serrata) and ginkgo (Ginkgo biloba), but because of the more immediate need to have a forest with an atmosphere appropriate for a shrine, a 150-year program was devised, whereby fastgrowing trees\u2014most notably Hinoki cypress (Chamaecyparis obtusa), Japanese cryptomeria (Cryptomeria japonica), and two species of pine (Pinus densiflora and P. thunbergii)\u2014provided at least the appearance of a chinju no mori before the slower-growing, broadleaf species gradually took over.7 Even with fifty of the 150-year program remaining, the Meiji Jingu\u00af forest has begun to feel old. It attracts ten million visitors every year, with three million coming in the three days after the New Year to pray. Wishes for the coming year are written on wooden tablets called ema, and these are left at the foot of a camphor tree (Cinnamomum camphora), which is believed to transmit the wishes to the deified emperor and empress. Visitors repay the trees with affection and respect, and the shrine's tree-viewing etiquette is rigidly adhered to. As soon as visitors pass through the ceremonial wooden torii (gate) they are in sacred space, and a quiet, respectful demeanor is assumed. No one leaves the paths to walk on the forest floor; no one picks leaves or seeds from a tree; and no 26 Arnoldia 77\/4 \u2022 May 2020 one removes anything from the forest\u2014even fallen leaves are left on the ground. Fabled Trees of Tokyo The desire to connect with Japan's spiritual and mythic past is accompanied by a strong feeling of connectedness with the nation's history. As a result, a number of Tokyo parks and gardens with old trees that have witnessed the city's history unfold have been given status as national monuments and historical landmarks. One of the most popular of Tokyo's historic landmark parks is the Institute for Nature Study, a 49-acre (20-hectare) forest that doubles as a research facility and a green oasis for the people of Tokyo. The Institute for Nature Study was once the feudal estate of the Matsudaira, a samurai clan related to the shogun (the military ruler of Japan). The star attraction is the Fabled Pine, an enormous Japanese black pine (Pinus thunbergii) that was part of the Matsudaira garden in the early 1600s. People typically bow before the old tree, which provides a living link to this emblematic era of Japanese culture\u2014a symbol of cultural continuity. The Fabled Pine also offers a reassuring example of resilience. The old tree's never-say-die spirit was underscored when the second-oldest tree in the Institute for Nature Study forest, a Japanese black pine known as the Ancient Pine, was toppled by a typhoon in October 2019. The fact that the Fabled Pine survived when even its venerable neighbor succumbed has only added to its mystique. Indeed, many of the visitors who stream up to the tree every day would be aware that the tree had survived a long list of disasters that began with the Great Fire of Meireki, in 1657. That fire burned 70 percent of the city and took over one hundred thousand lives\u2014far more destructive and deadly than the Great Fire of London, nine years later. Earthquakes were also a constant threat. Major quakes hit the city A ceremonial torii leads into the Meiji Jingu\u00af forest. STEPHEN SCHNEIDER Old Trees in Japan 27 in 1703, 1855, and 1894; then in 1923, the biggest of them all, the Great Kanto\u00af Earthquake, flattened most of the city. During World War II, Tokyo was spared the horror of the atomic bomb, but the city's trees were decimated by the relentless American firebombing.8 The chances of any tree surviving this litany of disasters is illustrated by the fact that of the twenty thousand trees in Shinjuku Gyoen, one of Tokyo's largest parks, only two, a 150-yearold magnolia (Magnolia denudata) and a 400-year-old zelkova (Zelkova serrata), are over one hundred years old. The zelkova\u2014the star attraction\u2014is showing signs of age. Its trunk was severed about ten feet from the ground, and new branches poke through a protective coat that was wrapped around the trunk to nurse the tree back to health. On face value, a tree that needs to be nursed back to health is an unlikely symbol of resilience, but as J. W. T. Mason has explained, according to Japanese tradition, great age and \"special hardihood\" are evidence of a tree's \"vital powers.\"9 The broken trunk and protective coat emphasize the battles the tree has fought, and give heart to residents of Tokyo that they can cope with the stresses and strains of their daily commute, long working hours, or, if they are young, looming exams. Survivor Trees of Hiroshima Never was Japanese resilience tested more than in the aftermath of the atomic bomb dropped on Hiroshima, in 1945. The bomb blast and resulting fires killed 140,000 people and destroyed all but a few buildings within an approximately 1.2-mile (2-kilometer) radius of the hypocenter. Survivors then began experiencing radiation sickness, resulting in death from cancer. At first it seemed as though the city's trees were following a similar trajectory. Most were instantly torn out of the ground or had their trunks snapped in half. The few trees left A prominent zelkova (Zelkova serrata) at Shinjuku Gyoen is more than four hundred years old. GLENN MOORE 28 Arnoldia 77\/4 \u2022 May 2020 standing were seared by a blast of heat so intense that a streetcar over a half mile (900 meters) from the hypocenter was completely oxidized. As one city administrator put it, the bomb had reduced the city to \"an ashen coloured wasteland bereft of all green.\"10 The fear was that nothing would grow in the radiation-affected soil for seventy-five years. Not surprisingly, residents were resigned to abandoning the city, when, almost miraculously, green shoots began emerging from some of the blackened, charred branches. A few trees, so burned and broken that they had no viable branches left, somehow managed to sprout new shoots out of their blackened stumps. A weeping willow (Salix babylonica), merely 0.2 miles (370 meters) from the hypocenter, was completely felled by the blast but managed to send up new shoots directly from its roots. In all, 170 trees regrew after the blast. Hibakusha (people who survived the bomb) have given testimony that the resilience shown by the hibaku jumoku (survivor trees) helped convince them that life could return to the city. Akio Nishikori was a second grader when the bomb fell. \"We were told nothing would grow for seventy-five years,\" he recalled. \"However, trees put out new shoots! Everyone was really moved to see the green leaves. These trees were the first to encourage humans [to rebuild.]\"11 In 1946, governor Kusunose Tsunei enlisted six community representatives to help him formulate a plan for restoring the city. As a city administrator explained, the consensus was not to \"create everything anew.\" Rather, the aim was to restore the \"social functions, culture, and traditions that had existed in the Hiroshima communities before the bombing.\"12 This meant many things. Hiroshima Castle, flattened by the bomb, was rebuilt. Hondo\u00afri, the city's ornate shopping arcade, was restored to its former glory. But most importantly, Kusunose's panel was adamant that \"many trees should be replanted in the city.\"13 It was no small task to grow trees in soil that had been burned by a nuclear blast and that was laced with rubble and debris, but today Hiroshima is a green city. Trees growing in parks and along rivers and roads give Hiroshima the look and feel it had before the war. The survivor trees provided living links to that prewar period, and the city was effectively rebuilt around them. Commemorative plaques were installed, and the trees have been preserved and tended into old age, even in cases when it might have been more convenient to remove them. A fine example of this respect is an old ginkgo (Ginkgo biloba) that, before the bomb was dropped, stood on the grounds of the Hosenbo Temple. The temple was levelled by the blast, and the head priest and his family were killed. The ginkgo had branches torn off and was badly burned, but it survived. When rebuilding began in 1994, the priests realized that the ginkgo, now a very large tree, would have to be removed to accommodate the architectural plans. Not willing to cut down a tree that had displayed such courage, they asked the architect to alter the plans, so that the temple could be built around it, preserving the tree as a symbol of resilience and continuity. Walking among Old Trees In 1982, Tomohide Akiyama, director of the Japanese Ministry of Forestry and Fisheries, coined a new term: shinrin-yoku (forest bathing.) 14 In fact, although it was a new word, the idea was connected to the very old Japanese notion that being among trees was good for health. In time the idea would become mainstream, with books written about how to get the most out of forest bathing and with one thousand government-accredited Official Recreation Forests now including shinrin-yoku trails. But when Akiyama coined the term, in 1982, he was also responding to a growing unease about the shift from agrarian to urban lifestyles, and the stress of modern life. By the 1980s, 80 percent of the Japanese population was concentrated in cities, seemingly far removed from nature. Long commutes, even longer workdays, and the constant pressure of not making a mistake would lead to Japan becoming recognized as the most sleep-deprived country on earth. There is even a Japanese word\u2014karoshi\u2014for the concept of death by overwork.15 Initially, there was no scientific basis for Akiyama's assertion that \"being in the forest makes our bodies healthy,\" but there was a sense that the idea was at least plausible.16 Chiba Old Trees in Japan 29 University horticulture professor Yoshifumi Miyazaki, who would later conduct studies on the value of shinrin-yoku, described this prevailing sentiment: \"The practice of walking slowly through the woods, in no hurry,\" made \"intuitive\" sense to the Japanese.17 This intuition was partly rooted in knowing how much better it was to smell pine trees instead of car fumes, or to hear birds instead of harsh city noises, but at a spiritual level, it was also connected to the deep traditions associated with trees. As Shinto Studies professor Sadasumi Motegi put it, shrine forests and parks with old trees \"are places that remind one of distant, ancient times. This is where the voices of the gods (kamigami) sound in your ears. This is where our ancestors lived, humbly, in harmony with nature.\"18 In 1990, in a study funded by the Japanese national broadcaster NHK, Miyazaki set out to test whether the spiritual benefits of reconnecting with nature were matched by medical benefits. To that end, he monitored the effects of walking through a forest on stress hormone levels in the human body. The findings were promising but inconclusive. Subsequently, he received a large government-funded grant allowing him to conduct more detailed studies. These studies have shown that there are not only emotional benefits from spending time in a forest but also measurable physiological benefits. For instance, office workers with stress-related high blood pressure had their levels lowered after spending six hours in an old growth forest. But the truly remarkable thing was that those with low blood pressure had their levels raised.19 In effect, forest bathing, or shinrin-yoku, restores the balance that is so hard to achieve in modern life. While science has supported the effectiveness of shinrin-yoku, the spiritual element involved has made it harder to explain in scientific terms how it works. As Miyazaki conceded in 2018, \"we need to do more research.\" What is known, however, is that it works best in an unspoiled forest setting, pristine enough for moss to grow freely, and where old trees live. Moreover, shinrin-yoku requires intentionality to work GLENN MOORE GLENN MOORE A weeping willow (Salix babylonica, left) was among the 170 trees that survived the atomic bombing of Hiroshima, as was a ginkgo (Ginkgo biloba) at the Hosenbo Temple. correctly. As Qing Li, the chairman of the Japanese Society of Forest Medicine, cautioned, \"This is not exercise or hiking, or jogging. It is simply being in nature, connecting with it through our sense of sight, hearing taste, smell and touch.\"20 In other words, forest bathers should proceed at the same sedate pace and with the same quiet, respectful attitude as when they visit a shrine forest. Bridging Past and Present Japanese city parks all at least aspire to a natural, unspoiled look, and many contain astoundingly old trees. This design intention is evident, even to a traveler, looking for cherry blossoms. Flower beds, a staple of parks in Western cities, are rare, and any lawns come with rules prohibiting ball games, music, or other activities that would shatter the serenity. Jogging is rarely permitted. When the Fabled Pine at the Institute for Nature Study, in Tokyo, was first planted under the auspices of the Matsudaira clan, more than four hundred years ago, the residents of the estate could never have imagined modern cities like Tokyo or Hiroshima. However, one thing they would find comfortingly familiar is the sustained relationship with trees. Shrine trees are still sacred. And old trees are still revered for their resilience, and they still provide a bridge back to the past\u2014indeed, even to the Matsudaira clan itself. As visitors at the Institute for Nature Study walk along the narrow path that wends through the trees, around ponds and over creeks, the city Shinrin-yoku paths wind through the landscape at the Institute of Nature Study in Tokyo. GLENN MOORE seems a million miles away. It was as if the park was designed specifically with shinrin-yoku in mind. The older trees are not, as in many parks, \"features,\" standing unnaturally apart from the rest of the plants. Instead they rise through a bed of saplings and bushes. Moss covers everything, underscoring the sense that the trees are growing in a pure, natural environment. Although Tokyo has changed so much over the last four hundred years, the Matsudaira clan would surely recognize the thinking behind shinrin-yoku, namely that spending time with trees is a life-giving activity. Endnotes 1 Hearn, L. 1904. Kwaidan: Stories and studies of strange things in Japan. Boston: Houghton Mifflin, 139-141. 2 Ishikura, Y. 2019, April 3. Hanami a reminder of life's fleeting nature. Japan Times. Retrieved from https:\/\/www.japantimes.co.jp\/opinion\/2019\/04\/03\/ commentary\/japan-commentary\/hanami-reminderlifes- fleeting-nature 3 Shirane, H. 2013. Japan and the culture of the four seasons: Nature, literature, and the arts. New York: Columbia University Press, 128. Note: According to Shirane, farmers believed that trees had emotions, and a tree would scream or groan when it was cut. 4 De Mente, B. 2018. Japan: A guide to traditions, customs, and etiquette. Tokyo: Tuttle, 10. (Original work published as Kata: The key to understanding & dealing with the Japanese, 2003). 5 Yamakage, M. 2006. The essence of Shinto: Japan's spiritual heart. Tokyo: Kodansha International, 11. See also: Hardacre, H. 2017. Shinto: A history. New York: Oxford University Press, 16. 6 Rots, A. 2017. Shinto, nature and ideology in contemporary Japan: Making sacred forests. London: Bloomsbury, 85. 7 Matsui, T. 1996. Meiji Shrine: An early old-growth forest creation in Tokyo. Restoration and Management Notes, 14(1): 46-52. See also: Saigusa, N. 2005, November. A 150 year-project: Meiji Shrine forest in central Tokyo. Japan For Sustainability Newsletter, no. 39. Retrieved from https:\/\/www.japanfs.org\/en\/ news\/archives\/news_id027807.html. 8 Cheng, S. and McBride, J. 2014. Restoration of the urban forests of Tokyo and Hiroshima following World War II. In K. Tidball and M. Krasny (Eds.), Greening in the red zone: Disaster, resilience and community greening. New York: Springer Books, 225. Note: Ironically, because records were destroyed in the fire-bombing, the exact number of trees lost from parks and gardens is unavailable. However, it is known that over sixty thousand street trees were destroyed in the war. 9 Mason, J. W. T. 2002. The meaning of Shinto. Victoria, Canada: Trafford, 75. (Original work published 1935.) 10 City of Hiroshima. (n. d.) A history of Hiroshima's greenery. Retrieved February 6, 2020, from http:\/\/www. city.hiroshima.lg.jp\/www\/contents\/1274090206341\/ index.html 11 Green Legacy Hiroshima. (n. d.) An introductory video [Video file in Japanese, with English subtitles]. Retrieved from http:\/\/glh.unitar.org\/ 12 Hiroshima for Global Peace Plan Joint Project Executive Committee. 2015 Hiroshima's path to reconstruction. Hiroshima: Rijo Printing, 24. 13 Kosakai, Y. 2009. Hiroshima peace reader. Hiroshima: Hiroshima Peace Culture Foundation, 18-19. 14 Miyazaki, Y. 2018. Shinrin-yoku: The Japanese way of forest bathing for health and relaxation. London: Hachette, 10-24. 15 Hoffman, M. 2018, September 8. Japan loses sleep over a variety of modern-day issues. Japan Times. Retrieved from https:\/\/www.japantimes.co.jp\/news\/2018\/09\/08\/ national\/media-national\/japan-loses-sleep-varietymodern- day-issues. Note: A 2016 survey ranked Japan last out of one hundred nations in terms of hours slept per night. Half of Japanese workers get less than six hours sleep a night. 16 Hendy, A. 2018, June 4. The call of the wild: Forest bathing and urban greening. The Japan Journal Online. Retrieved from https:\/\/www.japanjournal.jp\/science\/ environment\/pt20180604165729.html 17 Miyazaki, 9. 18 Rots, 85. 19 Miyazaki, 24. Note: for an example of a recent study on the effect of shinrin-yoku, see: Song, I. and Miyazaki, Y. 2017. Sustained effects of a forest therapy program on the blood pressure of office workers. Urban Forestry and Urban Greening, 27: 246-252. 20 Li, Q. 2018. Into the forest: How trees can help you find health and happiness. London: Penguin Random House, 12. Acknowledgments The authors would like to thank the staff at the Management Office of the Shinjuku Gyoen for their help and advice. Glenn Moore taught for many years at the University of Melbourne. He now works for the Japanese national broadcaster Nippon Hoso Kyokai (NHK). Cassandra Atherton is an associate professor in Writing and Literature at Deakin University. Old Trees in Japan 31"},{"has_event_date":0,"type":"arnoldia","title":"Each Year in the Forest: Spring","article_sequence":5,"start_page":32,"end_page":40,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25705","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d2708126.jpg","volume":77,"issue_number":4,"year":2020,"series":null,"season":null,"authors":"Hipp, Andrew L.","article_content":"Each Year in the Forest: Spring Andrew L. Hipp Illustrated by Rachel D. Davis I I spend the last weeks of each February brushing leaves aside, anticipating spring shoots. The first I find is often false mermaid, born beneath the maple litter. Its three-lobed leaves fold over themselves like the fingers of a glove. The fleshy cotyledons, newly hatched from the seed, are embedded a millimeter or two below the surface of the soil, soft and green on their inner surfaces, roughened on the backs where the clay and sand cling. The roots are spidery and translucent, barely a fifth of the height of the plant. The long petiole is ghostly white at the base and striated with elongating cells. It gradually darkens to a pale green just below the leaf blade, while the rest of the plant unrolls at the base. Once the false mermaid is out, I know all hell is about to break loose in the understory. Soon wild leeks prickle from the soil's surface, pale at the tips with crimson sheaths. Cut-leaved toothwort arches as it extracts its inflorescence from the soil. Its leaves are feathery and purple. Spring beauty reclines beneath the duff accumulated at the bases of the tree trunks or matted on the forest floor, fragile white stems spreading into green, strap-like, delicious leaves that Colony of Wild Leeks Forest: Spring 33 have not yet become bitter. Some of the spring beauty grow so far in search of light that their stems become threadlike. Bullet-shaped mayapple shoot-tips cluster at the bases of the oaks, each tip a half-inch high, clothed in white scales, like a goblin's fingertips. Wood violet rhizomes squeeze out infinitesimal soft green leaves. Rain buries the rhizomes in floods of soil and washes them downslope. Still they continue to grow. Mosses green up on decomposing logs and on soil that was sterilized by the burning of cut buckthorn in previous years. The mosses form a bed for flowering plant seedlings and a barrier between spongy, rotting wood and the desiccating air.* Mats of wavy starburst moss bristle with sporophytes, capsules at the tips of the filaments popping open to release a little dust storm of spores when I brush them. The spores float off and settle onto nearby plants and logs. Sporophytes wearing slender hoods emerge from a tussock of baby tooth moss, resembling bristles on a hairbrush. The leaves are as thin as the pages of a Bible, with the smallest possible teeth on the margins. Over the next few days, the capsules begin to swell and bend, and soon they are nearly perpendicular to the sporophyte stalks. The hoods split along the side. Within two or three weeks, they will be fully reflexed. They produce spores before most flowering plants in the woods start exchanging pollen. Evergreen leaves that have been working all through the winter give way. White bear sedge sends up blue-green shoots from broad-leaved, leathery rosettes. Pennsylvania sedge produces slender tillers, and even the leaves that overwintered brighten up. Winter leaves of white avens and strawberry and hepatica continue photosynthesizing as they pass the baton to the young leaves, which emerge as wrinkled as newborns, readying themselves to become next year's evergreen foliage. False Mermaid Baby Tooth Moss Bristling with Sporophytes 34 Arnoldia 77\/4 \u2022 May 2020 Infant soil centipedes curl on red oak chips that are decomposing in beds of earthworm castings. Chorus frogs sing and then retreat as the temperature swings. Spring peepers awaken and accompany me on my morning walks with strident, individual squeaks. I follow them into a weedy marsh, and suddenly I am in a fog of peepers blasting away, chorus frogs creaking behind them, but not a one to be seen. As I walk away, their calls attenuate, and in two hundred feet I no longer hear them. It is this way with peepers: a person could go through an entire spring within a quarter mile of a pond of peepers awakening and wooing in springtime and never know they were there. II That was the first week of March. Soon, wild garlic sprouts from gravelly roadsides and trail edges, slender as grass. Scales loosen on the flower buds of silver maples growing along city streets and creeks. The scale margins whiten with hairs, then the buds open and spill out a handful of stamens or dark-red, tentacular paired styles. Hazelnut catkins descend and are at first stiff, then looser a few days later, bracts cupped like umbrellas over the anthers balled up inside. Then they relax just a bit more and the anthers open. I cannot resist tapping the catkins growing together on a shrub, making them wobble like rows of prayer wheels and release clouds of pollen. Male woodcocks skate through early March mornings, peenting in openings in the grasslands before soaring overhead, making a distinctive kissing sound White Bear Sedge Pennsylvania Sedge Forest: Spring 35 when they reach the top of their aerial dance. On my bike ride into work at the Morton Arboretum, in the western suburbs of Chicago, I often hear the woodcocks spinning over the arboretum's easternmost marshes and fields as I unlock the gate. It is still dark out, and they are flying high overhead before they drop onto gravel roads, openings in the marshes, or mown fields around the cultivated collections. When I hear one calling from the ground, I will sometimes wait for the flight upward, then race to where he was. Almost invariably he drops down too far away, and I don't find him. Once last spring I succeeded in seeing one drop back and resume his dance. He barely lifted his chin when he called. After each \"peent,\" he paused and did a head-nodding shuffle forward and then backward before calling again. He seemed to wait for a response each time, shuffling as though in anticipation of the next call, a restless suitor. He called about five times before growing silent and then abruptly flying off to circle overhead. We'll have a few weeks of dancing woodcocks before they grow quiet and a portion of the flock moves farther north. We'll see them again on their way back through in the fall. Flocks of juncos buzz and pop in the shrubs, tails flashing as they whip back and forth over the trail. Last year's stump puffballs show up crushed against logs, and expired earthstar fungi nest in the wood chips. White ice fills ephemeral pools like congealed clouds on days when the temperatures rise to 50\u00b0F (10\u00b0C). The ice then melts outward from the maples and elms that perforate the pools, until the water is wide open, with only a glaze of clear ice returning on late-March mornings when temperatures dip below freezing. False mermaid, now a few inches tall, bunches up in openings in the oak leaves. Bluebirds perch on the lateral branches of bur oaks and scan the thawing turf for insects. Eastern phoebes return. One day, near the end of the month, I hear the protracted bubbling song of the winter wren. It stops for a few seconds, then starts again, five seconds of a complex line. The song twists around tree trunks and lichen-covered branches that were knocked to the ground by winter storms. I follow it and, if I am lucky, find the wren picking its way among mosses and scraps of soft wood, from one end of a rotten log to the other. A flock of American robins spreads out across the forest floor, solitary birds flipping leaves over one by one, looking for millipedes and pillbugs. It will be completely silent except for an occasional chuckle from the robins and the sound of leaves rustling, which might be the wind's doing if it were not the robins'. III Early April stammers as temperatures drop. This is the lull before the pandemonium of spring wildflowers. False mermaid is widespread, but not thick anywhere. It bolts, overtops the mottled sheets of oak and sugar maple leaves, and spreads across the bare soil of ephemeral watercourses. A week later, it carpets the woods. Spring beauty sprawls beneath the oaks, flower petals streaked with pink. Bloodroot flowers emerge, stalks wrapped in the solitary leaf. Rain a few 36 Arnoldia 77\/4 \u2022 May 2020 days later knocks their petals to the ground. Jewelweed cotyledons pop out on bare upland soil and floodplains, each the size of a nickel, fleshy and bitter. The lavender flowers of hepatica arise beside its light-green, rubbery new leaves, often at the bases of oaks where the plants are protected and where they can soak up rain that flows down furrows of the bark. The white flowers of false rue anemone pool in colonies scattered throughout the woods. One morning, on my bike ride into work, I find the field sparrows have started claiming territory. Their bouncing song rings through the woods for a minute before I reach a field embedded in the woods. Chipping sparrows trill and harvest insects from the swelling oak buds. Tree swallows patrol the birdhouses. Ruby-crowned kinglets flit in the lower areas of the woods, moving continuously, singing an uncontainable song that breaks open and spills through the leaves around me. Chorus frogs and spring peepers are exuberant and everywhere. I park my bike and walk in, and when I pause to list the birds I've been hearing, the first tick of the season crawls across my notebook. Mourning cloak butterflies come out from beneath panels of tree bark where they have slept out the winter. Bumblebees and painted ladies cross the trail. Then a late-season snowfall buries the wildflowers. Mayapples huddle in bunches against the snow, like passengers waiting for the bus in a blizzard, False Rue Anemone Jack-in-the-Pulpit Forest: Spring 37 leaves tucked tightly under their chins. Spring beauty in full flower reclines against a log where it is protected from the drifts. Wild leeks and Virginia bluebells are rigid, frozen in mid-expansion, figures in a wax museum. Then the next morning it is 60\u00b0F (15\u00b0C), and the snow melts away. By mid to late April, Dutchman's breeches forms puddles of foliage on slopes and disturbed trail margins. Its flowers school above the leaves. When the plants first emerged in early March, I hardly noticed them, flower buds condensed like frog eggs on the translucent scapes. Now, the white flowers mature from bottom to top, petals stretched back into deep spurs, stigmas arched at the snout. Jack-in-the-pulpit spears upward through the foliage before it grows tall enough to spread its wings. Soon it sends up a slender, fleshy inflorescence axis packed densely with pistillate or staminate flowers that I only see by carefully peeling back the hood. Wild ginger leaves appear at the tips of the rhizomes, folded over one another as they emerge, light green and hairy among trampled dead maple leaves. They spread open as soon as their blades are free of the earth, then lie back to sop up the sun. Jewelweed cotyledons I noticed in early March give way to scallop-margined foliage. And leaves begin to come out on the trees, unfurling like wet handkerchiefs on sugar maple seedlings and dripping from the tips of the hackberry branches. False mermaid has grown lanky. One day I notice its three diminutive petals, about two-thirds as long as the green sepals that alternate between them. They form a crown around three or six stamens, tipped with yellow anthers, and two or three prickly ovaries. The flowers, like the plants themselves, are easy to miss if you are not watching closely, and I sometimes miss their opening. If I have been particularly inattentive, the ovaries may already be swelling by the time I first see the flowers. The first plant I watched for in February does not flower until the spring ephemerals\u2014toothwort, spring beauty, Virginia bluebells\u2014 are already in full bloom. IV Yellow-rumped warblers appear near the middle or end of April with little warning. I typically hear them before I see them, singing from high in the canopy, and I struggle to remember whose song it is until I see the warblers stalking among the branches or catching flies midair. I will perhaps have already noticed blue-gray gnatcatchers bizzing and wheezing along the tree branches. Soon after, black-throated green warblers show up in the neighborhood, singing in the highest tree branches on our street as they warm up in the mornings. I know we are in the thick of warbler migration when I hear the lazy \"bee-buzz\" of the blue-winged warblers coming from trees along the edges of fields. Ovenbirds call insistently from the shrubbiest areas of the woods. Black-and-white warblers squeak in the midstory like rusted bearings. Last year, one struck an herbarium window at the arboretum and lay stunned. His eyes closed slowly, and he rolled onto his side on the window ledge. I reached out to retrieve him, but he flipped over and flew off between the branches of the European beech that shades the window. 38 Arnoldia 77\/4 \u2022 May 2020 At about the time that the yellow warblers and common yellowthroats start singing, wood thrushes return to a stand of closely planted spruces embedded in the arboretum's East Woods. The stand is low, with intermittently running water and a thick undergrowth of wood nettles that by this time is tall enough to sting my knees. This habitat seems to be just right for the wood thrushes. In the afternoons, orioles will be tearing at catkins in the tops of the red oaks and piping their hearts out. One evening, American toads begin droning from the marshes. Their song spills out into the adjacent forest. There are a few more weeks left in spring, but we are at the turning point to summer. Wildflowers flood the woods, running in sheets across the fallen oak leaves and overtopping the spring foliage that has carpeted the woods in the past two weeks. Spring ephemerals have peaked and begun to fruit, as they race to complete their entire annual life cycle on the sunny forest floor before the leaves are fully out on the trees. Cut-leaved toothwort, which flowers with petals the size of a child's incisors, produces siliques, slender capsules that crack open along the sides to release an abundance of small seeds. The flowers on Dutchman's breeches ripen to capsules. Rivers of Virginia bluebells flower, then the corollas fall off, leaving the capillary style ringed at its base with swelling hard nutlets. The first flowers of wild ginger open beneath the foliage, a pelage of long hairs combed over the backsides of the calyx, purple sepals tipping backwards. Anthers dangle from tiny flowers on male plants of early meadow-rue, and the females' flowers are frosted with stigmas. Rue anemone forms beds of beautiful, full-faced white blooms, some doubled so you might take them for cultivars. Glaucous branches of blue cohosh twist like dancers. Capsules swell thick as bullets on bloodroot. As the canopy begins to close, the wildflowers of late spring take over. Wild geraniums form lavender seas. The trilliums flower: first bloody butcher with purple petals arching upward, then large white trillium, and then nodding American Toad Forest: Spring 39 trillium, petals stretching out from between the sepals. Flowers dangle like bells in the leaf axils of Solomon's seal and hairy Solomon's seal; their leaves resemble those of the false Solomon's seal and starry Solomon's plume, but the flowers of those species form bouquets at the tips of the stem. The understory burns with wild hyacinth. V Everything that was brightest and most beautiful in mid-May is overrun by the end of the month, as wild lettuce reaches to my knees and orchard grass stretches out along the road through the arboretum. The false mermaid I found the first week of March is yellowing and flattened like seaweed against a boulder, pouring its last into the nutlets ripening at its apex. The forest floor is a bed of jewelweed. Yellowing leaves of white trout lily and variegated leaves of toothwort and wild leek stand out in the darkening understory beside the last flowers of false rue anemone. The first flowers of great waterleaf open as the hairy, spiderlike inflorescence branches unroll atop the plants. Bloodroot leaves swell to the size of my hand with fingers fully outstretched and lay back to absorb what sunlight they can through the closing canopy. Maple and elm seeds rain down overnight, clogging the gutters. Mayapple flowers become the lights of the woods, shining from beneath their great green umbrellas. For a week or so, I can hear golden-winged and black-and-white warblers, northern parulas, black-throated greens, all passing through, alongside the birds of summer: pewees and great-crested flycatchers, kingbirds, phoebes, gnatcatchers and red-eyed vireos, tanagers, ovenbirds, buntings, wood thrushes, and orioles. An olive-sided flycatcher calls an insistent \"quick three beers!\" Mosquitoes become pesky in the evenings. Spring peepers grow silent. American toads drone on. Wild Ginger PLANTS REFERENCED The onslaught of spring has come to a close, that time when I see each plant from all sides and keep thinking, what will happen tomorrow? Because for a few weeks, everything is happening at once. No one could catch it all in one year. A person needs year after year in, ideally, a single forest to get the sequence straight. * For more on the ecology, beauty, and importance of mosses, read Robin Wall Kimmerer's magnificent Gathering Moss: A Natural and Cultural History of Mosses (2003, Oregon State University Press). 40 Arnoldia 77\/4 \u2022 May 2020 Geranium maculatum - wild geranium Geum canadense - white avens Hepatica acutiloba, H. americana - hepatica Hydrophyllum appendiculatum - great waterleaf Impatiens capensis, I. pallida - jewelweed Lactuca spp. - wild lettuces Laportea canadensis - wood nettle Maianthemum racemosum - false Solomon's seal Maianthemum stellatum - starry Solomon's plume Mertensia virginica - Virginia bluebells Plagiomnium cuspidatum - baby tooth moss Podophyllum peltatum - mayapple Polygonatum biflorum - Solomon's seal Polygonatum pubescens - hairy Solomon's seal Rhamnus cathartica - buckthorn Quercus macrocarpa - bur oak Quercus rubra - red oak Sanguinaria canadensis - bloodroot Thalictrum dioicum - early meadow-rue Thalictrum thalictroides - rue anemone Trillium flexipes - nodding trillium Trillium grandiflorum - large white trillium Trillium recurvatum - bloody butcher Viola sororia - wood violet Acer saccharinum - silver maple Acer saccharum - sugar maple Allium canadense - wild garlic Allium tricoccum - wild leek Arisaema triphyllum - Jack-in-the-pulpit Asarum canadense - wild ginger Atrichum altecristatum - wavy starburst moss Camassia scilloides - wild hyacinth Cardamine concatenata - cut-leaved toothwort Carex albursina - white bear sedge Carex pensylvanica - Pennsylvania sedge Caulophyllum thalictroides - blue cohosh Celtis occidentalis - hackberry Claytonia virginica - spring beauty Corylus americana - hazelnut; you may also have C. cornuta in your area Dactylis glomerata - orchard grass Dicentra cucullaria - Dutchman's breeches Enemion biternatum - false rue anemone Erythronium albidum - white trout-lily Floerkea proserpinaca - false mermaid Fragaria virginiana - strawberry; you may also encounter F. vesca as a common species in your area Andrew Hipp is the senior scientist in plant systematics and herbarium director at the Morton Arboretum in Lisle, Illinois. He conducts research on the origins and implications of plant diversity, with a focus on oaks, sedges, phylogenetic ecology, and trait evolution. You can read about his research at http:\/\/systematics.mortonarb.org and follow his natural history blog at https:\/\/botanistsfieldnotes.com. Rachel Davis is an independent visual artist in the Chicago area. She works at the interface of natural science, abstract painting, printmaking, and textiles, integrating the formal and empirical elements of the natural world in her work. You can see more of her work at https:\/\/ artbumble.com and follow her on Instagram: @art_bumble."},{"has_event_date":0,"type":"arnoldia","title":"How to See Urban Plants","article_sequence":6,"start_page":41,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25706","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270816a.jpg","volume":77,"issue_number":4,"year":2020,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"\"Sight is a faculty; seeing, an art,\" the environmentalist George Perkins Marsh wrote in his 1864 treatise Man and Nature. \"I know no more important practical lessons in this earthly life of ours\u2014which, to the wise man, is a school from the cradle to the grave\u2014than those relating to the employment of the sense of vision in the study of nature.\" Botanical field guides, which became increasingly popular around the turn of the twentieth century, aimed to support the art of seeing. Their authors promised to transform overlooked spaces into landscapes filled with interest. One of the first successful field guides for the northeastern United States was Frances Theodora Parsons's How to Know the Wild Flowers, which was originally published in 1893. In the introduction, Parsons attested that \"even a bowing acquaintance with flowers \u2026 causes the monotony of a drive through an ordinarily uninteresting country to be forgotten in the diversion of noting the wayside flowers, and counting a hundred different species where formerly less than a dozen would have been detected.\" Yet an individual field guide necessarily elevates certain plants over others. After all, the guide must be sized for a pocket or, perhaps more How to See Urban Plants Jonathan Damery 42 Arnoldia 77\/4 \u2022 May 2020 realistically, a backpack. One of the most provocative field guides to appear in recent years is Peter Del Tredici's Wild Urban Plants of the Northeast (Cornell University Press), which presents an expansive vision for which plants, not to mention which landscapes, are worthy of being seen. The second edition was published this spring, adding forty-five plants to the twohundred- plus included in the 2010 edition. Del Tredici, who is an emeritus research scientist at the Arnold Arboretum, has provided, among other things, photographic documentation of the overlooked plants that inhabit overlooked urban places. In cities like Boston, New York, and Philadelphia, the plants are often so common that they are inconspicuous, ubiquitous but unseen. Del Tredici shows riverbank grape (Vitis riparia) cascading from powerlines in an alleyway and prostrate knotweed (Polygonum aviculare) trapping cigarette butts on a sidewalk. These are common scenes that urban commuters and pedestrians often pass without a second thought. Almost half of the species that are newly added in the second edition are North American natives, including familiar trees like the black walnut (Juglans nigra). These additions suggest the haziness inherent to determining whether something should be included or excluded from any field guide\u2014even one, like Del Tredici's, that is emphatically inclusive. Del Tredici describes the black walnut as an ornamental shade tree from a bygone era. Its spread into neighboring lots may seem unremarkable; the trees are almost too normal to be noted. The same goes for the green ash (Fraxinus pennsylvanica), which was another new addition. Deciding which plants should be featured in a field guide necessarily requires parameters. Parsons, in 1893, described her intention of omitting plants that were \"so common as to be generally known\" and \"so inconspicuous as generally Previous page: horseweed (Erigeron canadensis) and dandelion (Taraxacum officinale). Above: tufted lovegrass (Eragrostis pectinacea) and prostrate knotweed (Polygonum aviculare). Wild Urban Plants 43 to escape notice.\" Del Tredici, meanwhile, draws lines pertaining to the definition of the term urban and, by extension, a distinction between the spontaneous and the cultivated. The most fascinating photographs in the book are those that show the plants within their urban milieu. After all, as Del Tredici writes in the introduction, \"it is the context in which the plant is growing\u2014not the plant itself\u2014that makes it a weed.\" Within these landscapes, the plants often appear uncontained; they have an agency unto themselves. Road markings are a frequent motif in the photographs, and Del Tredici captures the omnipresent dandelion (Taraxacum officinale) sandwiched between a left turn lane and the oncoming traffic, as though the plants were moving in flagrant disregard of the yellow centerlines. He shows an American elm (Ulmus americana) flattening itself against a chain-link fence, in Hartford, Connecticut, where it has been hacked back repeatedly. The photographs document a changing landscape, one caught in an ongoing state of becoming. The book includes more than one thousand of Del Tredici's photographs, and in the opening pages, he notes that many were taken on family trips and errands. He acknowledges his family's patience with \"sudden stops on the side of the road\" for \"yet another \u2018weed' picture.\" In this sense, the photographs are remarkable in that they document not merely the fact of the plants but also the fact of stopping for them\u2014the fact of pulling over to the side of the road, as other cars zipped past, and hopping onto the shoulder to actually observe the plants up close. The book, in other words, is a testament to the necessary art of seeing. Jonathan Damery is the editor of Arnoldia. Riverbank grape (Vitis riparia) crosses over a Detroit alleyway, and maintenance crews would surely love to remove this American elm (Ulmus americana) in Hartford."},{"has_event_date":0,"type":"arnoldia","title":"Spring is the New Fall","article_sequence":7,"start_page":44,"end_page":45,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25707","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270856d.jpg","volume":77,"issue_number":4,"year":2020,"series":null,"season":null,"authors":"Schoonderwoerd, Kristel","article_content":"When we welcome new foliage in the spring, we must also bid adieu to another set of structures that have adorned trees throughout the winter: the bud scales. That's right, May is the perfect time for bud-scale peeping, and there is no finer tree to start with than the shagbark hickory (Carya ovata), a Massachusetts native. With its fingerlike leaflets pointing towards the sky, emerging from a skirt of magenta scales, the spring shoot of a shagbark hickory is reminiscent of an alien that has recently taken up flamenco dancing. A large grove of shagbark hickories (accession 12907) can be observed near the Centre Street entrance to the Arnold Arboretum, and for an observant spring enthusiast, the spectacular display is likely to turn an easily overlooked piece of botany into a pressing question: What are these pink structures? Buds scales are best known for their winter protective role. Trees repeatedly develop new organs (leaves) throughout their lifetimes, but this mode of continuous development can be challenging in a temperate climate. In the words of John Muir: \"Consider what centuries of storms have fallen upon [trees] since they were first planted,\u2014hail to break the tender seedlings; lightning, to scorch and shatter; snow, winds, and avalanches, to crush and overwhelm,\u2014 while the manifest result of all this wild storm-culture is the glorious perfection we behold.\" Bud scales are one item in a long list of adaptations that make this glorious perfection possible, and not an insignificant one. Bud scales envelop the sites where new leaves are initiated. Newly formed miniature leaves, waiting for spring, are thus provided with a sheltered space for their earliest development. As the leaves mature within the bud scales, their familiar form, comprising a leaf blade, a leaf stalk, and a leaf base (the attachment point of the leaf to the stem), begins to appear. To understand what bud scales truly are, in addition to what they do, we have to follow this closely coordinated chain of leaf development even further back to when the scales themselves were first formed. Bud scales are, in fact, leaves\u2014 modified leaves, never meant to capture light over the growing season. When a tree builds a bud scale, it makes a leaf with just a long and thin version of a leaf base and none of the other components. We know this because vein patterns in the base of certain photosynthetic leaves are similar to those in bud scales. In addition, if you look at a lot of bud scales, and you are lucky, then you might find a happy accident where a leaf has ended up half bud scale and half foliage leaf, indicating the relatedness between the two forms. Moreover, we know that bud scales and photosynthetic leaves are initiated in the same pattern, from the same cell clusters. New bud scales appear relatively early in the season for shagbark hickories. In fact, if you examine the tender stems that emerge with the fresh leaves in the spring, you can already see the very earliest, minute instances of the new bud scales\u2014including the pink flamenco dresses of the following year. This way, the growing tips are not only protected during winter but are never once exposed through the four seasons. It follows that the conspicuous pink phase in early May is but the swan song of the shagbark hickory's bud scales. After many months of passive sheltering, the scales start to rapidly expand and change color in the spring. The reddish color may well point towards a continued protective role. These blushes are caused by anthocyanins, the very same compounds that color leaves red or purple in the fall. The currently most-favored hypothesis states that anthocyanins function as a sunscreen, protecting delicate structures\u2014new, growing leaves in this case\u2014from excess sunlight. Perhaps the spring metamorphosis observed in the shagbark bud scales is a final act to guard the small but rapidly expanding leaves, until the leaves can grow and function without outside protection and the scales can drop away, no longer needed, and make way for the next cohort. Kristel Schoonderwoerd is a doctoral candidate in the Friedman Lab at the Arnold Arboretum. Spring is the New Fall Kristel Schoonderwoerd"},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25702","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d270b727.jpg","title":"2020-77-4","volume":77,"issue_number":4,"year":2020,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"A Cottage Flora","article_sequence":1,"start_page":2,"end_page":5,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25673","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070bb6f.jpg","volume":77,"issue_number":3,"year":2020,"series":null,"season":null,"authors":"Meholic, Cat","article_content":"A Cottage Flora Cat Meholic I n the fall of 2015, I moved into a small cottage on the back of the Granogue Estate, a sprawling property in northern Delaware. When my parents first visited the estate, I could tell they were nervous. As we drove the milelong driveway, the road got worse and worse. I tried to understand their apprehension; their daughter would be living alone in the last cottage on a woodland edge. I only had one small dog at the time, a feisty, unintimidating Yorkshire terrier. As we drove past cornfields and open meadows, and descended into the valley, all I saw in my new home was a childhood dream come true. My parents admitted that the old stone house, covered in white plaster, was charming. MEHOLIC, C. 2020. A COTTAGE FLORA. ARNOLDIA, 77(3): 2?5 A young woodland is located on one side of my cottage, and the Brandywine Creek runs directly behind it. The light reflects off the creek into the house in unusual patterns, and on winter evenings, geese fly to the river, passing so low that I can hear the whispering of their wings. I often think of the Sand County Almanac and search for signs of life, as Aldo Leopold described, in every season: the mergansers that appear like clockwork in February, the tracks left from a battle of fox and rabbit in the snow, and the beaks of trillium (Trillium spp.) poking up in the spring. I am captivated by my surroundings. These woods have provided me with years of comfort and continuously pique my curiosity. Each copse is unique, but A Cottage Flora 3 most contain a mix of old tulip poplars (Liriodendron tulipifera), American beech (Fagus grandifolia), and hickories, primarily shagbarks (Carya ovata). The house itself is two stories, built into the hillside. When the house was inhabited by a farmer, his wife, and eight children, the downstairs kitchen was an open structure for the livestock to bed down at night. Today the cottage (or \"studio,\" as the residents of the estate call it) is stacked with horticulture and botany books. These books are mostly gifts from my mentors, colleagues, and friends, and the bulk came from the lifetime collection of my graduate advisor, John Frett. His collection was so extensive that I turned an open staircase into a makeshift bookshelf to house them. The generous windows overflow with plants. In this ideal setting, I have completed a thesis, adopted two more dogs, started a nonprofit called Women in Horticulture, and begun a checklist for the flora of the property. Ir?n?e du Pont Sr. established the Granogue Estate and relocated his family to the property in 1923, when he was president of the DuPont company. The main house is now the residence of Barbara and Ir?n?e du Pont Jr. Much of the 505-acre property is actively farmed for corn, soy, hay, and dairy production, but large sections of forest and meadow have been preserved. Although the estate is less than a twenty-minute drive from Trader Joe's, Target, and a shopping mall, the landscape feels like a rural oasis. In these woods and meadows, I have walked my dogs almost every day for four years. While holding two leashes, and with a third dog strapped to my waist, I scribble out notes in a Moleskine notebook tucked in my dog-walking fanny pack, recording the flora that I observe. Although I must be a comical sight to my neighbors, which happen to be mostly cows, this method has been effective. Upon returning home, I add additional details to the notebook, and on rainy days when I am not outside as much, I update my Excel spreadsheet. Over time this exercise has turned into a checklist that is extraordinarily simple. The list itself records just the scientific name of the plant, and in some cases the date I observed it. What started as a means of learning about the land I lived on quickly morphed into a love of the plant communities that inhabit there. Already this information has had small impacts on land use. One of my first successes on the property was when a trail-running race agreed to no longer use a path that was carved straight through a population of goldenseal (Hydrastis canadensis). When the trail was created, before a March event, I was disheartened that I would not be seeing the glorious little white flowers or finally observing the goldenseal fruit in a wild population. I travel regularly in this section of the woods, so this part of the population was the easiest for me to view while corralling my dogs along the path. I showed the damage to an ecologist friend, and she was also dismayed. Goldenseal is not exceptionally rare in the state, but this population is the largest one we had ever seen in northern Delaware. After several attempts, I succeeded in contacting the race director and did my best to communicate how the path impacted that population of goldenseal. This was after two seasons of the race using the path. Thankfully, the director was amenable to my recommendations, and the population is slowly rebounding from the disturbance. On another occasion, regular path maintenance was endangering a small group of common moonseed (Menispermum canadense). In this case, the damage to the population could not be avoided as this section of the property needs to be accessible by vehicles and a horsedrawn buggy. The woodland edge and the moonseed population had been slowly encroaching for years. To help preserve the genetic diversity of this population, whole plants were given to two botanical gardens: Mt. Cuba Center and Natural Lands' public garden, Stoneleigh. After path edges were mown back, the moonseed has surprisingly rebounded from its roots. The estate has also been a great resource for educational exercises. Mt. Cuba Center is located just 7.4 miles from the Granogue Estate and is a regional resource for everything related Facing page: An unnamed creek on the Granogue Estate flows into the Brandywine Creek. ALL PHOTOS BY THE AUTHOR 4 Arnoldia 77\/3 ? February 2020 tage is situated across a gravel driveway from a field for dairy cows. I am lucky that this field is used for breeding the next generation of bovines and not full dairy production. In the spring I have wrestled a newborn calf in the snow to tuck it back under the fence with its mother. For two years, however, I watched as cows defecated directly into a water source that drained into the Brandywine. Our analysis found that the small corridor of trees that served as a riparian buffer drastically reduced the amount of pollutants entering the Brandywine, confirming the ecological value of the plant populations that were included on my checklist. I am still hesitant to jump in the water downstream after an extreme rain event, but at least the impacts are much less than I anticipated. to native plants. I know the precise distance because I drove to Mt. Cuba every day for years as their plant recorder and assistant curator. Over the years I have had the pleasure of sharing the botanical treasures of Granogue with colleagues and friends, including those at Mt. Cuba. (I must admit, at this stage in my career, the terms \"colleague\" and \"friend\" are often synonymous.) Colleagues from Mt. Cuba were impressed by the extent of a large population of showy orchis (Galearis spectabilis) at Granogue, and two interns were sent to Granogue to do a population estimate and record associated taxa. Research on native orchids has been at the forefront of Mt. Cuba's research initiatives, and data from Granogue were included in work by Adrienne Bozic, the orchid fellow at Mt. Cuba, who oversaw the development of an orchid inventory for a large part of Delaware. I also worked with colleagues at the University of Delaware, where I completed my graduate work, to conduct an exploratory study on the impact an existing riparian buffer has on the water quality of the Brandywine Creek. My cot- My observations of the flora at Granogue also include comparisons to adjacent sites. The Brandywine Creek State Park is separated from Granogue by Thompsons Bridge Road. When crossing this road, the change in vegetation is apparent even to the most plant-blind of indi- A view of the Granogue Estate, with the Brandywine Creek State Park in the background and the author's cottage nestled into the tree line. A Cottage Flora 5 The state park has increased human usage, and the estate, meanwhile, has increased deer hunting, lowering herbivory pressure. Locals have also suggested that the presence of cows on the estate deters deer, which seems to be true, at least anecdotally. viduals. Although both sides have almost an identical canopy, the understory is drastically different. The state park has large swathes of Japanese barberry (Berberis thunbergii) that create six-foot walls on either side of the path. Porcelain berry (Ampelopsis brevipedunculata) covers defoliated trunks, and whole patches of forest have toppled due to stress, pests, and repeated wet summers. The Granogue side of the road has faced the same stressors, but the understory is more complex, which seems to add resilience to the existing canopy. The understory shrubs and trees include spicebush (Lindera benzoin), common witch-hazel (Hamamelis virginiana), American bladdernut (Staphylea trifolia), hornbeam (Carpinus caroliniana), American holly (Ilex opaca), flowering dogwood (Cornus florida), black cherry (Prunus serotina), pinxterbloom azalea (Rhododendron periclymenoides), swamp azalea (Rhododendron viscosum), smooth blackhaw (Viburnum prunifolium), and mountain laurel (Kalmia latifolia). The herbaceous layer is a fantastic mix of ferns, spring ephemerals, violets, sedges, and other native flora, including wild ginger (Asarum canadense), zigzag goldenrod (Solidago flexicaulis), white wood aster (Eurybia divaricata), numerous species of trillium (Trillium spp.), and yellow jewelweed (Impatiens pallida). Although most of these taxa are not considered rare, it is striking to see the contrast in population members between the state park on one side of Thompsons Bridge Road and the estate on the other. This difference in taxa has captured my curiosity. Both properties were historically logged and then fragmented into farm fields. The hills are steep and rocky along the Brandywine, with the iconic \"blue rocks\" that our Minor League Baseball team is named after, and these rocky slopes were often too difficult to use for crops but were moderately successful for grazing sheep or goats. Old stone walls, characteristic of New England and the northern Mid-Atlantic, cut through sections of the existing forest, acknowledging this past. Despite these similarities, certain site characteristics provide at least a partial explanation for the floristic differences between the properties: As stewards of your own properties or those publicly shared--neighborhood parks and even urban wilds--it is important to understand your land as thoroughly as possible, and part of this is to understand the plant and animal communities that are present. I'm reminded of a quote from Aldo Leopold: \"We abuse land because we regard it as a commodity belonging to us. When we see land as a community to which we belong, we may begin to use it with love and respect.\" From the time we are born, we learn to \"love and respect\" our human communities, but we must teach ourselves to do the same for our natural communities that surround us. As an observer of plant communities, I have seen how sharing the knowledge of these communities changes the way in which humans treat them. A checklist provides the basis for assessments of biological productivity, ecosystem classifications, conservation decisions, and documentation of spatial or temporal changes over time. Without a basic checklist none of this would be possible. Every effort towards a better understanding of our natural world counts. Without someone observing and documenting the plants at Granogue, projects involving the path modifications and subsequent off-site plant preservation would not have been possible. Even these small projects have had a positive impact beyond the Granogue boundaries. I might seem na?ve or romantic, but I firmly believe if more nature enthusiasts observed their surroundings more closely and acted on what they were seeing, the impacts would be magnified in meaningful ways. I hope that the trend of encouraging citizen scientists continues to expand until we roll our eyes at how everyone now calls themselves a citizen scientist. What better citizen could we hope for? Cat Meholic is the curatorial horticulturist at Ambler Arboretum of Temple University. "},{"has_event_date":0,"type":"arnoldia","title":"Beyond the Trees: An Herbaceous Shift at the Arnold Arboretum","article_sequence":2,"start_page":6,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25675","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070816d.jpg","volume":77,"issue_number":3,"year":2020,"series":null,"season":null,"authors":"Keegan, Brendan","article_content":"KEEGAN, B. 2020. BEYOND THE TREES: AN HERBACEOUS SHIFT AT THE ARNOLD ARBORETUM. ARNOLDIA, 77(3): 6?13 Beyond the Trees: An Herbaceous Shift at the Arnold Arboretum Brendan Keegan W hile walking down Oak Path last October, I was drawn to a glint of yellow among the dappled shades of green. I carefully stepped through the tall grasses and clumps of wood aster (Eurybia divaricata), and found, to my surprise, a blue-stemmed goldenrod (Solidago caesia) in bloom. A few bumblebees busily gathered pollen in the afternoon light, hanging on the plant's characteristically staggered flowers that ascend like golden rungs on a ladder. Although blue-stemmed goldenrod is a common native plant, I had never seen one growing in the middle of the oak collection. Despite providing ideal habitat, with the high canopy allowing ample sun, the collection is typically mowed, which kills young perennials long before their autumn blooms. In 2019, however, much of the oak collection was purposefully left fallow to encourage the growth of herbaceous species. The lack of mowing resulted in an abundance of common violets (Viola sororia) in the spring followed by blue and white asters (Symphyotrichum and Eurybia), several species of goldenrod (Solidago), and, presumably, some very pleased bumblebees in the fall. Unlike the accessioned trees (each lovingly labeled, monitored, and cared for), the wild populations of herbaceous plants, such as the bluestemmed goldenrod, often exist in anonymity. A new appreciation for the herbaceous understory's benefits to the woody collection and to the urban ecosystem, however, is redefining the Arboretum's approach to these plants. The horticulture team has, among other things, begun leaving areas fallow to encourage spontaneous populations. Staff have also begun collecting and propagating herbaceous plants for landscape renovations, and these plants have been introduced as ground cover in the formal collections. An herbaceous shift is underway. A Naturalistic Legacy Although the Arnold Arboretum is famous for its history of plant exploration and its collection of temperate woody species gathered from around the world, it is perhaps less well known that the landscape itself was originally intended as an homage to New England. Both Frederick Law Olmsted, the landscape architect who designed the Arboretum, and Charles Sprague Sargent, the Arboretum's founding director, guided this aesthetic, drawing on shared horticultural principles and a desire to mimic the region's natural ecosystems. Although the resulting naturalistic aesthetic is taken for granted these days, it is easy to overlook how innovative and even radical the Arboretum's landscape was considered at the time. By the late 1800s, city planners across the United States were beginning to think about the importance of green space for the public. The industrial revolution had brought the masses into the urban core, where there was little escape from toil in factories. Wealthy homeowners, on the other hand, developed their gardens in the fashion of the day, filling greenhouses with imported tropical plants and their yards with various exotic specimens. In many cities, newly built public parks reflected the taste of the wealthy, with formal gardens and annual beds taking pride of place. Olmsted's vision for the Arboretum, and for the larger parkway of which the Arboretum is one part, could not have been more different. He planned for the winding roads to be bordered by layers of natural-looking shrubs and trees in order to invoke feelings of rural country lanes. Olmsted preferred woods and natural areas, interspersed with sweeping lawns that were designed to be maintained by cows and sheep. This preference, however, didn't preclude the use of non-native plants. Olmsted outlined this Facing page: Kent Field provides a colorful showcase of herbaceous-layer restoration and management at the Arnold Arboretum. ALL PHOTOS BY JONATHAN DAMERY UNLESS NOTED ARNOLD ARBORETUM ARCHIVES 8 Arnoldia 77\/3 ? February 2020 Local photographer Thomas Marr captured the herbaceous diversity along Bussey Brook in 1908. idea in an 1888 letter to the editor, in Garden and Forest, a publication established and overseen by Sargent: \"May we not (as artists) think that there are places with us in which a landscape composition might be given a touch of grace, delicacy and fineness by the blending into a body of low, native tree foliage that of the Tamarisk or the Oleaster ... ?\" Fortunately, Olmsted had a more than willing partner in Sargent. Known for his passion for woody plant collection and discovery, Sargent formed the cast in which the Arboretum was molded. Many of the horticultural decisions he employed at the Arboretum could also be seen at his Brookline estate, Holm Lea. Although Sargent rarely opened Holm Lea to visitors, those granted access marveled at his natural- istic integration of native and non-native trees, shrubs, and herbaceous plants. Wilhem Miller, a writer for Country Life, praised the estate, in 1903, as an example of \"`natural' landscapegardening.\" Miller observed that \"while there are few places that are more cosmopolitan--so far as kinds of plants are concerned--the visitor will search in vain for anything that sounds a discordant note.\" Similarly, Nathaniel Greene, a journalist writing for the New England Magazine, in 1908, observed that \"Professor Sargent wants nothing on his place that is not harmonious with a New England landscape.\" The combination of Olmsted's landscape philosophy and Sargent's horticultural decisions resulted in an Arboretum that combines plants in a naturalistic way. Greene noted that this Herbaceous Shift 9 combination, as demonstrated at Holm Lea, differed from the approach used in parks in other cities, which \"tend to look alike, because they use chiefly tender bedding-plants, which are costly, ephemeral, loud, garish.\" A fitting, albeit melancholic, example of Sargent's lasting sympathies towards naturalistic plantings was the location of his memorial service held on June 9, 1927. Rather than celebrating his legacy near the Hunnewell Building where he spent much time, or under one of the species named in his honor, the event was held in one of his favorite locations along Bussey Brook \"where white oaks, a white pine, and a white beech, all native to American soil, overhung the temporary platform.\" The Spontaneous Flora While Olmsted designed the Arboretum's bucolic setting and Sargent filled it with plants from around the world, the responsibility of documenting the landscape's herbaceous species fell to staff working on the grounds. In the early years, this role was filled by Ernest Jesse Palmer, a self-taught botanist from southwestern Missouri. Palmer came to the attention of Sargent around 1901 after he mailed Sargent fruits from several species of hawthorn (Crataegus), a group of plants that Sargent was beginning to research intensely. The pair corresponded for decades, with Palmer regularly collecting plant material for the Arboretum, before he finally accepted a position as a botanist collector and moved to Boston in 1921. Palmer soon began to inventory and study the spontaneous herbaceous flora at the Arboretum. Fortunately, he assiduously documented his observations, with the vast majority of the 2,235 spontaneous plant vouchers in the Arboretum's herbarium attributed to his name. He compiled his findings in an article titled \"The Spontaneous Flora of the Arnold Arboretum,\" first published in a 1930 issue of the Journal of the Arnold Arboretum. Palmer's \"Spontaneous Flora\" remains a valuable benchmark for studying the diversity of wild plants in the Arboretum landscape, and the wider region. Unsurprisingly, the current wild plant composition is much different than Palmer's records describe. In his day, there was greater overall herbaceous plant diversity. Palmer noted \"bits of open meadow\" between the young trees in many of the collections, \"where the grass and other undergrowth is cut at infrequent intervals.\" Many plants flourished among what were then saplings in the hawthorn collection, including nine species of aster and goldenrod. Included among them were the \"pretty violet rays and yellows discs\" of the flax-leaved aster (now classified as Ionactis linariifolia), which were \"especially abundant and showy.\" These days, flax-leaved aster is a rare sight among the hawthorns. Other parts of the Arboretum, however, seem unchanged. Palmer observed, in the 1920s, that \"Elderberry ... Joe Pye Weed ... Poison Hemlock ... New England Aster ... and New York Aster are most conspicuous\" along the steep banks of Bussey Brook, a spot where they continue to thrive. Somewhat amusingly, he also noted that the \"curious little parasitic Love-vine or Dodder twines its amber evanescent stems about some of [these] upright herbs.\" Despite periodic assault by a century's worth of horticulturists intent on its removal, a healthy population of dodder (Cuscuta gronovii) persists in that exact location to this day. Palmer's wild plant baseline also proved invaluable for follow-up surveys documenting the change of spontaneous species through time. At least nine species from his 1930 publication no longer grow in the Arboretum, and four of those are now endangered in Massachusetts. Of the six species of orchid that once grew in the landscape, only pink lady slipper (Cypripedium acaule) has been recently documented. Ten additional herbaceous species are on the state watch list, and their status in the Arboretum is undetermined. Ironically, Palmer, who worked without the benefit of historical documentation to guide his observations, lamented that many plants which \"might have been found ... [are] now gone forever from the Arboretum\" as a result of physical alterations ongoing in his day. Cultural preferences likely played a role in changing species abundance. Lawns were increasingly viewed as more favorable than unkempt meadows, and they allowed greater mobility throughout the collection. Although horse-drawn sickle mowers were the tool of choice by the early 1900s (and were used by Sargent's land managers at Holm Lea), frequent mowing would not occur until the Arboretum 10 Arnoldia 77\/3 ? February 2020 Jerusalem artichoke (Helianthus tuberosus) and blue-stemmed goldenrod (Solidago caesia) attract summer pollinators. acquired gas-powered machinery in the midst of labor shortages during World War II: a tractor, with an attachment for hay cutting, in 1945, and two lawn mowers in 1946. Mowing also provided a cost-effective and reliable tool for keeping down unwanted vegetation, which became especially important as invasive plant populations increased. The asters and goldenrods that Palmer described on Peter's Hill were soon jostled by Oriental bittersweet (Celastrus orbiculatus) and black swallowwort (Cynanchum louiseae), just two of many invasive species that harm the valuable woody specimens they grow among and on. While frequent mowing can mitigate these problems, it also kills the native grasses and forbs that once beautified the collections, depressing overall biodiversity. Mowing has also resulted in mechanical damage to accessioned plants and contributed to soil compaction collection-wide. Although no comprehensive bulletin of spontaneous plants has been published since Palmer's last edition of \"Spontaneous Flora,\" in 1945, staff and researchers continue to assess the status of herbaceous plants and other spontaneous species. Peter Green updated Palmer's list, in 1962, when he published \"Herbaceous Aliens in the Arnold Arboretum\" in Arnoldia. Leslie Mehrhoff, the herbarium curator at the University of Connecticut, reviewed and updated Palmer's baseline while documenting changes in invasive plant populations beginning in 2008. Mehrhoff intended to publish a comprehensive update to Palmer's list, but the project ceased after his death in 2011. Around that time, Walter Kittredge, of the Harvard University Herbaria, completed a multi-year taxonomic project, verifying the identifications for the well-over two thousand herbarium specimens that constitute the Arboretum's spontaneous flora collection. These individuals have kept institutional knowledge alive by channeling Palmer's surveying efforts, allowing the Arboretum to keep track of changes to herbaceous species composition through the years. Shaping the Herbaceous Layer In 2017, new strategies for increasing the ecological value of the landscape began reshaping the herbaceous flora once again. These changes started when Andrew Gapinski, the Arboretum's head of horticulture, organized the first official Herbaceous Committee. The group, of which I was a founding member, was initially concerned with identifying areas in the landscape where mowing pressures could be reduced or eliminated, in an effort to limit mechanical and compaction-related damage to accessioned plants. Compacted soils had compounded after decades of running heavy, gas-powered equipment directly around and beneath the plants. A healthy herbaceous layer would allow for staff to leave \"no-mow\" zones, mitigating future compaction and reducing fossil fuel inputs. The committee also anticipated that, as shorterlived herbaceous species come and go, their Herbaceous Shift 11 Herbaceous plants provide visual texture throughout the seasons. decomposing roots would help to break up and enrich the compacted soil. The committee discussed ways to increase the herbaceous diversity and how these collections could subsequently be managed by horticultural and curatorial staff. Irina Kadis, former Arboretum curatorial assistant and native plant expert, was central to this effort. Her knowledge and cross-checking of Palmer's observations informed the committee's reintroductions. The committee also relied on her personal knowledge of natural lands to determine from where to source the plants. In order to maximize the ecological benefits, as well as to mimic the naturalistic setting which Palmer described, the committee agreed to only introduce plants native to Massachusetts. These plants would, ideally, be sourced or collected from wild populations, allowing the Arboretum to document provenance and conserve local ecotypes. This means that, for a widespread species like red columbine (Aquilegia candensis), which has a native distribution stretching from the East Coast to the eastern Great Plains, the Arboretum would only acquire locally sourced wild material, to protect our regional genetic diversity. Acquiring wild seed poses a significant hurdle and is an ambitious jump for an institution focused on woody plants. However, there is ample precedent for building the collections with plants from New England. In the 1870s, the Arboretum propagator, Jackson Dawson, used a horse and buggy to collect native shrubs and trees to fill the young Arboretum's nurseries. Dawson continued to make local collections throughout his more than forty-year career, but recent local efforts are the first to strategically add to the Arboretum's herbaceous understory. I helped organize collaborations with the Native Plant Trust, the Trustees of Reservations, and the Massachusetts Audubon Society, allowing staff to purchase or collect seed from beautiful natural areas close to home. These revised management strategies have resulted in prominent no-mow zones on Peters Hill and in the conifer, oak, birch, linden, and maple collections. The meadows on the face of Peters Hill are particularly notable. Although cool-season grasses comprise the majority of the spontaneous vegetation, large stands of common milkweed (Asclepias syriaca) and goldenrod are emerging. In addition, over two thousand butterfly milkweed (A. tuberosa) plants dot the hillside, propagated in the Dana Greenhouse from seeds collected on Martha's Vineyard and donated by the Polly Hill Arboretum. The meadow also provides habitat for a variety of insects, birds, and mammals. A pair of eastern bluebirds, spotted in the spring of 2019, were the first to successfully nest in the Arboretum's collection since 2006. In the Kent Field meadow, down the road from Sargent's chosen memorial site, masses of introduced mountain mint (Pycnanthemum muticum), common boneset (Eupatorium perfoliatum), butterfly milkweed (Asclepias 12 Arnoldia 77\/3 ? February 2020 Jim Papargiris, the Arboretum working foreperson, plants butterfly milkweed (Asclepias tuberosa) during a staff workday in 2018. tuberosa), little bluestem (Schizachyrium scoparium), great blue lobelia (Lobelia siphilitica) and golden Alexanders (Zizia aurea) provide food and shelter for wildlife throughout the seasons. In late summer, monarch butterfly caterpillars carefully munch on the milkweed, while a variety of butterflies and bumblebees cover the stands of mountain mint and purplestemmed aster (Symphyotrichum puniceum). In the evening, Kent Field is a prime spot to observe bats darting for insects, while great horned owls use the towering conifers to watch for unwary voles and mice. In addition to revitalizing meadows, the Arboretum is also actively collecting and purchasing herbaceous plants in response to other forms of landscape change. In 2018, a considerable number of beeches (Fagus) were removed from the Arboretum's collection, due to an outbreak of beech bark disease. This caused a massive shift in the growing conditions for the herbaceous layer, which went from deep shade to full sun essentially overnight. Expansive areas were laid open and bare. The following year, I coordinated the purchase of several thousand plugs of wild-sourced foxglove beardtongue (Penstemon digitalis), New York aster (Symphyotrichum novi-belgii), and other herbaceous species from the Native Plant Trust. Horticulturist Scott Phillips and I developed a plan to use these native plants for landscape restoration in the collection. Although most of the herbaceous plantings have occurred in naturalistic portions of the Arboretum landscape, the efforts have also filtered into the formally designed collections. Phillips led the acquisition of herbaceous species to fill mulch beds in the Bradley Rosaceous Collection. This project fulfills the original vision of the Bradley redesign, which began in Herbaceous Shift 13 The Peters Hill meadow provides habitat for nesting birds. 2007. The herbaceous plants, primarily native members of the rose family (Rosaceae), fill in the open expanses of mulched beds, complement accessioned species, mitigate weed problems, and reduce the future needs for herbicide applications. Above all else, the new plants are beautiful and highlight ways that green groundcovers can be simultaneously functional, attractive, and consistent with the overall vision of the collection. Constant Change Temperate woody plants will always be the cornerstone of the Arboretum's mission, but recent projects with herbaceous plants will hopefully continue to spread through the collection. These projects will be crucial for the Arboretum's efforts to develop a more resilient landscape, protecting our valuable woody plants while enhancing the ecological function of our urban habitat. Beyond that, these herbaceous plantings fit well with the intent of both Olmsted and Sargent, whose original designs stressed the importance of a naturalistic landscape. Looking to the past verifies that the only constant with plants in the Arboretum's landscape is change. As Palmer observed, herbaceous populations and entire species will continue to come and go, following the ebbs and flows of management, culture, and climatic changes. If the trend continues, perhaps herbaceous accessions will eventually lose their anonymity and become as beloved to visitors as the towering trees. In the meantime, you can be sure that at least one Arboretum demographic will enjoy them--bumblebees searching for just a few more blue-stemmed goldenrod flowers on a warm autumn afternoon. Brendan Keegan is a gardener at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"A Lily from the Valley","article_sequence":3,"start_page":14,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25674","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d0708128.jpg","volume":77,"issue_number":3,"year":2020,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"A Lily from the Valley Michael S. Dosmann T his story is of a flower and the man who ventured to the other side of the world, away from family and modern convenience, to collect it. The plant was not just an object of desire but one of such value that it would underwrite the most significant collecting expeditions of the day. Yet, its beauty almost betrayed the collector, nearly taking his reputation and his life. It is also a story of their redemption: the story of Ernest Henry Wilson and the regal lily (Lilium regale). When I first glimpsed regal lilies in the wild, in 2014, I was in northern Sichuan Province, China, to retell Wilson's story for CCTV's documentary, Chinese Wilson. I recall how gusts filled the air with sand, as well as a bright aroma from lilies, prompting me to simultaneously squint and sniff deeply. Ensembles of the glistening, trumpet-like blossoms dotted the gray cliffs above the Min River. The blooms were sometimes a half-dozen to a stalk, predominantly a clear white, with a purplish blush on the outside and yellow throats within. DOSMANN, M.S. 2020. A LILY FROM THE VALLEY. ARNOLDIA, 77(3): 14?25 I admired them from a vantage point on the narrow, rocky trail below. Most stems reached straight up to the sky, while others dangled out from the cliffs at near-ninety-degree angles. I was baffled by how they could defy gravity like that, with so little soil to cling to amidst the ever-blowing wind. Because of my plant collecting experience in China for the Arnold Arboretum, following in Wilson's footsteps, I had been asked to guide viewers for the documentary. The third and final episode highlighted Wilson's collection of Lilium regale and a rockslide that nearly ended his life not far from where we filmed that day. The episode was rounded out with narrations of Wilson's own descriptions of events. That part was easy. Wilson retold the story often, in numerous books and articles, with a dramatic flair that would have prompted Mark Twain's praise. Most accounts started with a rehearsed rhetorical question, as it did in \"Price of the Regal Lily,\" published in Country Gentlemen in October 1925: \"How many people know the E.H. Wilson and Lilium regale 15 The author (facing page) is photographed for a documentary that recounted Ernest Henry Wilson's harrowing collection of the regal lily (Lilium regale). The filmmakers orchestrated a reenactment of the mule train that was important to Wilson's retellings of the story. size of a mule's hoof?\" He then would respond, \"Frankly I do not know with mathematical exactness, but as I lay on the ground and more than forty of these animals stepped over my prostrate form the hoof seemed enormous, blotting out my view of the heavens.\" How is that for an opening line? The explorer went on to richly describe the dusty \"rude land\" south of Songpan where his \"royal lady\" grew: \"That such a rare jewel should have its home in so remote and arid a region of the world seemed like a joke on Nature's part.\" The disaster occurred on September 4, 1910, while Wilson was on his fourth expedition to China. \"Dysentery in a mild form\" had prompted him to ride in the sedan chair, yet he noted that \"song was in [their] hearts\" for they were near Wenchuan and just north of Sichuan's capital, Chengdu, where good food and accommodation awaited. When the landslide struck, his chair was tossed to the river several hundred feet below. Errant boulders left the team scattered, and Wilson's right leg shattered in two places. Luckily, he never lost consciousness, and he instructed his team to use the camera tripod to splint his leg. It was then that PHOTOS, PAGE 14 BY KOU JIN, PAGE 15 BY MICHAEL S. DOSMANN the mule train approached. Because the path between the cliff face and the roaring torrents below was too skinny for them to turn around, the only choice was for Wilson to remain on the ground and watch as each and every mule stepped over. What followed was a hastened and painful three-day journey to Chengdu, with Wilson carried on an improvised stretcher constructed from the remnants of his chair. Doctors at the Friends Foreign Mission set his leg as best they could, but the possibility of amputation persisted for weeks due to nagging infection. In the end, however, his leg--now nearly an inch shorter than his left--was saved, as were the lilies. During Wilson's recuperation, members of his team dug up a quantity of bulbs, which followed Wilson back to Boston in the spring of 1911. Wilson was so proud of the introduction that, despite the near-death experience and life-long injury, he stated that the \"lily was worth it and more.\" In his 1925 monograph The Lilies of Eastern Asia, he went even further, proclaiming that \"in adding it to western gardens the discoverer would proudly rest his reputation with the Regal Lily.\" I concur, this lily is a gem. But ARNOLD ARBORETUM ARCHIVES 16 Arnoldia 77\/3 ? February 2020 Ernest Henry Wilson (right) and zoologist Walter Zappey rest along a footpath in central China in 1908. ger for their (and his) value to be realized? Or, was there something more to his statement-- did Wilson really believe his reputation was at stake and only redeemed by this lily? Wilson was responsible for introducing over a thousand plants to Western cultivation, including scores of horticultural prizes. The ghostly dove tree (Davidia involucrata) haunted his dreams on his first expedition for Veitch Nursery, and the yellow poppywort (Meconopsis integrifolia) was his muse for the second. He had also introduced his favorite shrub of all time--the beautybush (Kolkwitzia amabilis)-- and the paperbark maple (Acer griseum). Wilson considered the maple, whose namesake bark is loved by connoisseurs everywhere, Hubei's best. Perhaps these successes didn't register to him because another collector sent one dove tree seed to France before Wilson managed to collect his bundle, and the poppywort proved a bit too finicky to cultivate broadly. As for the other two woody plants, maybe it just took lon- Little is written about Wilson's state of mind during his days of exploration, and his own correspondence barely sheds light upon such things. (Personal letters to his wife, Nellie, were destroyed by the family after the couple's death in 1930.) His journal entries have hardly seen the light of day due to his near-indecipherable penmanship, but one entry stands out beyond others, written on September 3, 1910, the day before the landslide. Wilson described his stomach trouble, his inability to keep warm, and the terrible road conditions. He noted the abundance of regal lilies (known then as Lilium E.H. Wilson and the Regal Lily 17 myriophyllum) upon the cliffs and described how, earlier in the day, two members of his team stayed behind in Sian Sou Qiao to investigate the region's conifers and to secure bulbs. The final paragraph is the most profound. While a word or two still evade \"translation,\" Wilson wrote of being in the same area two and a half years before. It had rained then, too, and I can imagine the drudgery, even misery, of being ill, sopping wet, loaded down with supplies, and trudging along a dangerous road still days away from civilization and convenience. \"I little thought then I should ever return here!\" Wilson lamented. \"I am certainly getting very tired of the wandering life & long for the end to come. I seem never to have done anything other than wander wander through China!\" Between 1899 and 1911, Wilson spent almost eleven years wandering through China, despite having a wife and, eventually, a young daughter, Muriel, at home. He was tired of the explorer's life before he wrote this entry in 1910 and was reluctant to head back after returning from his second trip for James Veitch & Sons nursery in 1905. He was then working as a botanist at the Imperial Institute of Science in London and lived at Kew, just a short walk from the Royal Botanic Gardens' gate. But, the stubborn persistence of Arnold Arboretum director Charles Sprague Sargent (and his accomplice Ellen Willmott, who worked the local English angle) finally persuaded Wilson to return to China in 1907, for what he thought was a final time. Whereas his trips for Veitch were motivated more by profit than botany, his work for the Arnold Arboretum was a scientific endeavor, with value placed on the germplasm secured in seeds, cuttings, and plants, as well as on the collection of welldocumented herbarium vouchers and photographs. Sargent, however, had arranged for a certain procurement of bulbs, which would help subsidize the 1907 expedition. Wilson first met the regal lily in August of 1903 while traversing the Min River Valley; the following autumn he sent about three hundred bulbs to Veitch under collection number 1791. They arrived in England in the spring of 1905, flowered that summer, and were identified at Kew as Lilium myriophyllum, a species described by the French botanist Adrian Ren? Franchet in 1892. Much was made of the freeflowering plants, with Wilson writing about the collection that year in Flora and Sylva. In 1906, Curtis's Botanical Magazine profiled the new-to-cultivation species, complete with a beautiful illustration. By the close of 1906, Sargent not only secured Wilson as the Arboretum's collector in China but found a partner to share some of the financial burden: John K.M.L. Farquhar. The Scottish-born nurseryman had established R.&J. Farquhar & Co. in 1884. It became one of the most prominent horticultural businesses in America, operating out of Boston. On Christmas Eve of 1906, Sargent wrote to Farquhar, \"Since our conversation of the other day I have talked over the bulb business with Wilson and have reached the conclusion ... that for the species from western China, namely ... [L.] myriophyllum ... thirty-five cents a bulb would be a fair price, in view of the fact that these would have to be carried on men's backs for at least two hundred miles before water transportation is reached.\" Two days later, Farquhar accepted the proposal, signing a contract to receive two separate shipments of bulbs collected by Wilson, paying all freight costs and a steep price for each sound bulb delivered. In the winter of 1907, Wilson found himself back in China and in no time reassembled his team in Yichang, Hubei Province. The collecting was good--Wilson began to accumulate vouchers, photographs, and plant material (including two Acer griseum seedlings that still grow in the Arboretum's collection). His first batch of lilies was also coming along nicely. According to the Farquhar contract, Wilson was to collect from \"Central China\" (namely Hubei) ten thousand bulbs, mostly the strident orange Lilium henryi but also L. leucanthum var. chloraster and L. brownii, both creamy white. (A collection like this would be unthinkable to modern collectors, not just logistically but because it is wholly unethical to dig up bulbs like this.) For those, Farquhar would pay $0.25 each (about $7 today). In a letter to Sargent before the turn of the year, Wilson commented that he would meet the quota but was worried about the cost of freight due to the quantity and weight of the cases. Rather than balling each bulb in clay, as he had done previously for Veitch, he informed ARNOLD ARBORETUM ARCHIVES 18 Arnoldia 77\/3 ? February 2020 On February 2, 1909, Wilson photographed men in Yichang packing cases of lily bulbs for shipment. For this second shipment to Boston, all of the bulbs were balled in clay. Sargent that \"this year I intend to try packing in dry sand only. This method ought to succeed but I know I shall be broken up if it fails.\" On January 17, 1908, thirteen cases--containing eleven thousand bulbs in total--left Yichang, travelling by ship down the Yangtze for Shanghai, then to England, and eventually Boston. Wilson ended up compromising on the packing. The Lilium henryi were packed in sand; the other two species were balled in clay. \"This is an experiment tried on the grounds of economy in freight and packing cases,\" Wilson wrote in a letter to Farquhar on January 29. \"For if it succeeds both parties benefit. If it fails both suffer loss.\" Adjusting for inflation, the bounty would fetch a sum of about $77,000 today. Farquhar would have his bulbs, Sargent a subsidized expedition, and Wilson the satisfaction of another job well done. Wilson and his team departed Hubei that spring and headed west into Sichuan for the second part of what he thought was his final campaign. In late May and June of 1908, he saw his \"royal lady\" in bloom in the Min River Valley near Wenchuan and Maoxian and made multiple herbarium vouchers under number 1446. (These were later designated as type specimens for Lilium regale.) No doubt, he was gearing up for the next round of bulb collecting to occur that autumn. In August, Wilson received a letter from Sargent, sent April 25. The news was devastating. Sargent reported that of the six thousand or so bulbs of Lilium henryi, which were not balled in clay, only four to five hundred had survived. Although it appeared that those encased in clay fared better (at least the bulbs sent to Sargent), most cuttings, grafts, and seeds of tree species had also died. \"The loss of the bulbs, however is a secondary matter as that is only the loss of money,\" Sargent wrote. \"In the loss of cuttings and grafts of plants like Willows, Poplars and Elms, the matter is much more serious because we have not seeds of these and you are not likely to be in a region to obtain them again.\" Sargent added, \"We are all, of course, greatly E.H. Wilson and the Regal Lily 19 disappointed over the outcome of this consignment, but, as I said before, I feel absolutely sure that you did what you thought was best.\" After receiving this devastating message, Wilson responded, \"I need not enter into my feelings of bitter disappointment and vexation on mastering its contents. In slang language I was `knocked all of a heap.'\" He promised Sargent he would \"remedy the failure.\" On October 30, Sargent wrote to Wilson: \"If it is possible to make up the loss in Farquhar's Lily bulbs, I hope you will do so, as we counted on the profit from these bulbs to pay a considerable part of the expenses of the expedition.\" This time, instead of the long-about method of getting to Boston via Europe, the bulbs would be shipped to the West Coast and travel across the continent on the Canadian Pacific Railroad (the method that Farquhar used to transport bulbs from Japan). And they would all be encased in clay, regardless of the extra freight costs. According to Farquhar's contract, the second shipment of another ten thousand bubs from \"Western China\" (namely Sichuan) would be shipped out in February 1909. This colorful motley would comprise equal numbers of Lilium bakerianum, L. leucanthum, L. duchartrei, L. sutchuenense (a synonym of L. davidii), and, of course, the regal lily. For these, Farquhar would pay $0.35 for each sound bulb delivered to Boston (about $9.90 today). Wilson rallied to meet this and then some. He added a few L. lophophorum to the mix and, in a letter to Sargent on December 29, reported that he had secured a total of twenty thousand lily bulbs, all balled in clay. \"Last year's experiment in attempted economy has been enough!\" he wrote. When the bulbs left Yichang for Boston, on February 20, 1909, the thirty-two cases included over two thousand bulbs of regal lily. \"This collection is a large one, and has been got together at a great expenditure of energy, indeed, I hardly know how it has been obtained,\" Wilson wrote to Sargent on March 9. \"If the bulbs arrive safely Messrs. Farquhar should not complain of there being nearly 20 instead of 10,000.\" Wilson continued with a boastful reflection: \"It gives them, I make bold to say, the finest chance they will have of securing not only the largest collection of Lilies from Western China that has ever been made but the only large one that will be made for decades to come.\" Wilson had no intention of returning. Without a doubt, such a quantity of bulbs would satiate the enterprising nurseryman. The higher premium ($0.35 a bulb) would satisfy the Arboretum's chief as well, for it would amount to almost $200,000 today. And lastly, having rectified the previous year's failure, Wilson could wrap up his work in China and return to England and his family. He left Beijing in April via train, eventually taking the Trans-Siberian Railway across the expansive Russian landmass to Moscow. From there, he continued to the major cities of Europe, visiting nurseries, gardens, and herbaria along the way. By the middle of May, he reunited with his family in England and was soon looking at plants collected on earlier expeditions and reviewing the photographs that he took on the recent trip. Waiting for Wilson at Kew, however, was a letter from Sargent, dated May 24. Sargent began by addressing an issue that must have caused him--and Wilson--some consternation: the issue of other botanical explorers in China. \"Sometime ago you wrote me expressing regret that the opportunity had not been given you to remain longer in China. This I confess was a very great surprise to me for you had told me more than once that nothing would induce you to remain in China for more than two years.\" In 1905, Frank Meyer began to explore China on behalf of the United States Department of Agriculture (and the Arboretum, when he found woody species of interest). And in February of 1909, Sargent and Veitch Nursery jointly dispatched another Kew graduate, William Purdom, to pick up where Wilson was leaving off. While Wilson was eager to end the arduous work in China, he was also worried about his reputation and the prospect of being replaced. In the letter that Sargent referred to, dated March 9, 1909, Wilson discussed both Meyer and Purdom, and he admitted to \"a slight feeling of chagrin at being passed over so completely in favour of another and without a word of warning.\" He continued: \"It can be interpreted unfavourable on the work I have accomplished during the past two years. I merely mention this--I do not say I think it thus intended.\" ARNOLD ARBORETUM ARCHIVES 20 Arnoldia 77\/3 ? February 2020 Snow covered the hills south of Yichang, as Wilson prepared to leave China in 1909, for what he thought was the final time. Wilson took this photograph on January 21 of that year. And then, Sargent dropped the other shoe. In addition to the thirty-two cases shipped to Farquhar, another five (including three cases of bulbs and other plants for Sargent's private garden and friends) were shipped to the Arboretum. Not only had the smaller shipment \"arrived in the most unsatisfactory condition,\" Sargent wrote that the \"bulbs sent to me were in much worse condition than those of the previous shipping. I do not think there is life in one per cent. of them.\" As if Wilson couldn't realize the magnitude of the loss on his own, Sargent spelled it out: \"This is, of course, a serious matter for the Arboretum as it involves a loss of probably six or seven thousand dollars which there is now no way of making up.\" In a follow-up letter to Wilson on June 3, Sargent confirmed that Farquhar's bulbs suffered similarly. An annotated manifest noted that just 121 of the 2,182 regal lily bulbs were alive at the time of arrival. Despite careful packing, the bulbs rotted in the ship's cargo hold. On June 9, Wilson wrote to Sargent: \"The disastrous news you sent, re. the condition of shipments, is a severe blow to me.\" Wilson had spent two years of rigorous and dangerous work in China, away from his family and alone save the companionship of his Chinese team (which included Walter Zappey, who collected alongside Wilson for Harvard's Museum of Comparative Zoology). His own legacy's status loomed in his mind well before getting this latest news, and with this failure, Wilson likely felt his reputation would suffer. Perhaps recognizing Wilson's state, Sargent proposed that Wilson come to Boston that summer to work through the innumerable herbarium vouchers. Wilson--now unemployed and much in need of a salary--agreed, noting, \"It will also allow the `rounding off' of the expedition in a manner I hope completely to your satisfaction.\" Sargent still described the expedition as successful in a letter to Ellen Willmott on August 23, no doubt because of the photographs, vouch- ARNOLD ARBORETUM ARCHIVES E.H. Wilson and the Regal Lily 21 Wilson photographed the habitat of the regal lily on August 31, 1910, just a few days before the landslide. \"A typical view in upper Min Valley,\" Wilson later captioned the image, \"showing barren desolate nature of the country.\" tributed to the Arboretum's Chinese Exploration Fund in hopes of a few plants of their own. The Wilsons departed Boston for England in the winter of 1910. Nellie and Muriel remained with relatives while Wilson retraced his journey via train back to Beijing. ers, and germplasm that had, in fact, survived. However, noting that the bulb debacle had cost the Arboretum nearly $8,000 (about $225,000 today), Sargent reminded her that she needed to remit to him the sum of ?6.10.3 (about $1,000 today) for her subscription to Wilson's expedition over the past two years. That September, Wilson, his wife, and daughter sailed for Boston, and he was soon organizing his herbarium specimens and doing his best to properly identify those lacking names. Nothing documents the conversations that must have occurred between him and Sargent, but within a few months, Wilson was planning a fourth trip to China. How much of this was due to Sargent's coaxing and how much of it was Wilson's need for redemption, we do not know. It was likely a mixture of both. Wilson planned a yearlong trip to Sichuan, with a focus on conifers that had evaded him before. To subsidize the expedition, Farquhar would still pay $0.35 for each bulb, while other private sponsors con- After the landslide and after doctors reset Wilson's leg, a Canadian Pacific Railroad train from Vancouver arrived in Boston. It was April 20, 1911, and the shipment carried Farquhar's complete order of bulbs, including some six thousand of the regal lily. They were immediately placed on the ground at the nursery and covered with soil. That summer, they flowered with wanton abandon, producing copious seeds by October. In Farquhar's Autumn Catalog, bulbs were already selling for $1.50 apiece ($40 today). Farquhar's Garden Annual of January 1912 lauded the regal lily, particularly the flower's unoppressive, jasmine-like perfume, and pre- 22 Arnoldia 77\/3 ? February 2020 dicted it the Easter lily of the future. The Massachusetts Horticultural Society awarded it a Gold Medal, and a beautiful illustration graced the November cover of The Garden Magazine (the American publication, not to be confused with the journal of the Royal Horticultural Society). Further admiration for it and other lilies appeared in an article in the same issue, with Farquhar's advertisements promoting their near-exclusive corner on the market. Wilson--the one who brought the horticultural world the regal lily--saw his reputation climb with that of the plant. The species, profiled on page one of Farquhar's Garden Annual of 1913, was attributed to \"the indefatigable plant collector, Mr. E.H. Wilson,\" who had collected it \"in remote and hitherto unexplored regions.\" That June, Wilson set the taxonomic record straight in The Gardeners' Chronicle, distinguishing Lilium regale from L. myriophyllum, the regal lily's maiden moniker. In this short article, Wilson also told the tale about the bulbs' transport \"on men's backs and by riverway 2,000 miles across China\" while he \"accompanied them in a stretcher or on crutches.\" While not as colorful and descriptive as his future retellings, Wilson was finding his voice. He was certainly getting much practice; in the same year, he published A Naturalist in Western China, a two-volume set of narratives about his travels. Farquhar's field of regal lilies in Roslindale, barely one mile south of the Arboretum, was abundantly populated, drawing crowds each summer. The Horticultural Club of Boston-- founded in late 1911 with John Farquhar and Wilson as inaugural president and secretary, respectively--made special fieldtrips to visit and witnessed some fifty thousand lilies in bloom in 1914. An article in The Florists Exchange titled \"Hardy Flowers at Farquhar's in July\" commented (perhaps with some hyperbole) on the lilies' display in 1916, noting that \"as many as thirty-eight fully developed flowers have been counted from one bulb on one stem, and a four year bulb will carry six stems.\" It was a popular item for sale and was frequently advertised in all the magazines. Farquhar's sale prices barely dropped to $1.25 a bulb through the teens, though there was the occasional offer of bulbs for $0.90 each. John Farquhar died in 1921, but the nursery continued under new leadership. Over the next decade, other nurseries such as Wayside Gardens (in Mentor, Ohio) and Baums (in Knoxville, Tennessee) promoted their own regal lily stock. Despite predictions that the regal lily would displace the common Easter lily as a forced bulb, production challenges limited this endeavor. A 1921 \"Talk of the Trade\" article in Horticulture Magazine noted how bulbs had to be \"carried over a year in a pot without having the flowers cut,\" which was impractical for most growers. Furthermore, a 1926 United States Department of Agriculture bulletin described how the market became flooded with smaller and smaller bulbs of poorer quality as growers offloaded stock, raising speculation about the species' worthiness. When Farquhar's nursery published its 1929 Garden Annual, regal lily was no longer profiled on page one, but was bundled with the other hardy lilies towards the back. Bulbs sold for $0.75 apiece, a price that continued to drop during the first few years of the Great Depression. Wilson, along with his wife, died in a car accident in the autumn of 1930. In 1932, R. & J. Farquhar Co. Nurseries went bankrupt and was resurrected as Dedham Nurseries. During the liquidation sale of all nursery stock, regal lily bulbs sold for just $0.15 each. The regal lily still sold through the midtwentieth century but was no longer an exclusive object of desire. Gardeners can be trendy, and it was the post-war era, when modern breeding programs were seen as the source of new plants, not old-fashioned field expeditions from a bygone age. George Pride, writing in these pages in 1974, summed it up: \"Although the Regal Lily has been superseded in favor with many gardeners by the fine modern trumpet strains of lilies, there are still gardeners who cherish and grow Lilium regale in its pristine, true species form and consider it still one of the best of all lilies.\" Brent and Becky's Bulbs of Virginia, one of the most well-known purveyors of geophytes in North America, currently sells the regal lily for $3.30 each. Facing page: Wilson and the regal lily (Lilium regale) were both celebrated in magazines, catalogues, and newspapers. BIODIVERSITY HERITAGE LIBRARY 23 ARNOLD ARBORETUM ARCHIVES 24 Arnoldia 77\/3 ? February 2020 While the regal lily was never officially planted in the Arboretum collections during Wilson's lifetime, Wilson cultivated a stand near his home on South Street. The original charter for the Arnold Arboretum, signed on March 29, 1872, declared that the living collections \"shall contain, as far as is practicable, all the trees, shrubs, and herbaceous plants, either indigenous or exotic, which can be raised in the open air.\" Even though herbaceous plants were included, Sargent, knowing the charge was too ambitious, soon adjusted the scope to focus solely upon woody plants. His reasoning also related to the Arboretum's relationship with the Harvard Botanic Garden, in Cambridge, and to his own desire to create something unique within the university. The botanic garden possessed well-ordered beds of herbaceous plantings, and it is likely Sargent had no interest in competing with them. He would set out to monopolize woody plants instead. And thus, not even one of Wilson's wildcollected Lilium regale bulbs was accessioned at the Arnold Arboretum. In fact, regal lilies from China were first accessioned in the autumn of 2017. Xinfen Gao, a professor of botany at the Chengdu Institute of Biology, had collected seeds while doing fieldwork near Maoxian, along the Min River. To no surprise, plants grown at her house flowered freely every year and set copious seed. She provided some to Andrew Gapinski and me for the Arboretum's collections at the conclusion of our expedition to Sichuan in 2017. Over a hundred bulbs from this accession were planted in the collections last autumn. This isn't the first time the species was grown on Arboretum property, however. Numerous lilies, including this one, grew in E.H. Wilson and the Regal Lily 25 Wilson's personal garden, an Arboretum-owned house across from the then Bussey Institute on South Street. And, in the fall of 1963, Lilium regale was included in a lily demonstration plot established at the Arboretum's Case Estates, in Weston. With Lilium regale finally growing in the Arboretum's collections, I cannot help but ponder the persistent allure of the species. With dogged determination, Wilson pursued it for years, and the lily still draws others to the Min River Valley, including the whole entourage who worked on the CCTV documentary. Wilson noted the regal lily was limited to a fiftymile stretch along the Min River, where it was nonetheless common. And, despite his removal of nearly nine thousand bulbs between 1903 and 1910, the species still flourishes and is not considered endangered (though it probably deserves protection). In fact, a recent paper by Wu Zhu-Hua and colleagues reported surprisingly high genetic diversity and no bottlenecks among the populations that scatter the cliffs along the Min, Heishui, and Zagunao Rivers (all within the Min River Valley). It seems that those ever-blowing gusts play a role in the regal lily's lasting reign, for the researchers attribute the species' survival to long-distance pollen and seed dispersal. When I was there, with the lilies' fragrance blowing in the wind, something else was also in the air: a siren's song-- or rather a lily's song--to lure someone back again and again. Acknowledgments I thank Lisa Pearson for her assistance in deciphering Wilson's handwriting, as well as her, Jonathan Shaw, and Jonathan Damery for their constructive comments during this article's development. Bibliography Anon. November 4, 1905. Lilium myriophyllum. The Gardeners' Chronicle III, 38, pp. 328?329. Anon. September 9, 1916. Hardy flowers at Farquhar's in July. The Florists' Exchange, pp. 589?590. Anon. 1921. The talk of the trade. Horticulture, 34(3): 53?54. Franchet, M.A. 1892. Les lis de la Chine et du Thibet. Journal de Botanique, 6(17?18): 305?321. Griffiths, D. 1926. The regal lily. United States Department of Agriculture Bulletin No. 1459. Washington, DC. Grove, A. April 27, 1912. New or noteworthy plants: Lilium myriophyllum. The Gardeners' Chronicle III, 51, pp. 272?273. Herrington, A. 1912. Lilies from June to October. The Garden Magazine, 16(4):145?147. Horticultural Club of Boston, Minutes and Records, 1911?1919 (volume 1). Archives of the Arnold Arboretum, Harvard University Pride, G. 1974. Lilies and the Arnold Arboretum. Arnoldia, 34(3): 125?132. Sargent, C.S. Correspondence (series III). Charles Sprague Sargent (1841?1927) papers, Archives of the Arnold Arboretum, Harvard University. Wilson, E.H. Correspondence, 1899?1930 (series W.XIV). Ernest Henry Wilson (1876?1930) papers, 1896?1952, Archives of the Arnold Arboretum, Harvard University. Wilson, E.H. Fourth Expedition to China--the Second for the Arnold Arboretum (series W.V). Ernest Henry Wilson (1876?1930) papers, 1896?1952, Archives of the Arnold Arboretum, Harvard University. Wilson, E.H. 1905. New and little-known lilies. Flora et Sylva, 3: 328?330. Wilson, E.H. June 21, 1913. New or noteworthy plants: Lilium regale. The Gardeners' Chronicle III, 53, p. 416. Wilson, E.H. 1915. Consider the lilies. The Garden Magazine, 21(6): 283?286. Wilson, E.H. 1925. The lilies of Eastern Asia. London: Dulau & Company Ltd. Wilson, E.H. 1925. Price of the regal lily: A treasure wrested from forbidding Tibet. The Country Gentleman. 90(36): 11, 145. Wright, C.H. 1906. Lilium myriophyllum. Curtis's Botanical Magazine, 132: Tab. 8102 Wu, Z.H., Shi, J., Xi, M.L., Jiang, F.X., Deng, M.W., & Dayanandan, S. 2015. Inter-simple sequence repeat data reveals high genetic diversity in wild populations of the narrowly distributed endemic Lilium regale in the Minjiang River Valley of China. PloS one, 10(3). doi:10.1371\/ journal.pone.0118831 Wyman, D. 1964. Lilies in their order of bloom. Arnoldia, 24(10): 89?95. Michael S. Dosmann is keeper of the living collections at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Confronting Climate Change at an Urban Grassland: Preserving and Restoring the Grasslands at Green-Wood","article_sequence":4,"start_page":26,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25676","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d0708526.jpg","volume":77,"issue_number":3,"year":2020,"series":null,"season":null,"authors":"Rossi, Frank S.; Evans, Sara; Charap, Joseph","article_content":"Confronting Climate Change at an Urban Grassland: Preserving and Restoring the Grasslands at Green-Wood Joseph Charap, Sara Evans, and Frank S. Rossi A ccording to standard ecoregion mapping of North America, New York City falls squarely within the eastern broadleaf forest--an ecosystem characterized by an overstory of tall, broadleaf trees, like American beech (Fagus grandifolia) and white oak (Quercus alba). The on-the-ground reality, however, is made obvious in the air: a plane flying low into LaGuardia International Airport offers its passengers a view of the city's expansive network of yards and parks, roads and parking lots. After generations of urbanization, New York City's tree canopy represents a mere 21 percent of its land cover. A comparable percentage is, in fact, represented by mixed grassland vegetation--the turfgrass in public parks, golf courses, and soccer fields. The city's cemeteries also house a significant portion of that vegetation, totaling an area of more than five Central Parks. At 478 acres, the Green-Wood Cemetery, in Brooklyn, is an active cemetery, a National Historic Landmark, and a curated arboretum, with a diverse and well-established collection of trees. It is also one of the largest urban grasslands in New York City. Founded in 1838, on land that was once forest, GreenWood includes rolling hills and kettle ponds. This topography, reflective of the landscape's position on a terminal moraine, made it inhospitable for agriculture but an ideal location to site a new kind of cemetery. Green-Wood was among the first cemeteries (after Mount Auburn Cemetery, in Cambridge, Massachusetts, and Laurel Hill Cemetery, in Philadelphia, Pennsylvania) built in the United States during the rural cemetery movement, a period in the mid-nineteenth century in which concerns over disease and rapidly increasing urban populations compelled city planners to site new burial grounds in the nearby countryside, instead of inner-city churchyards. In addition to providing burial space, these romantic, natu- CHARAP, J., EVANS, S., AND ROSSI, F.S. 2020. CONFRONTING CLIMATE CHANGE AT AN URBAN GRASSLAND: PRESERVING AND RESTORING THE GRASSLANDS AT GREEN-WOOD. ARNOLDIA, 77(3): 26?31 Urban Grasslands 27 ralistic landscapes served as counterpoints to the bustle and tumult of cities, providing sites for passive recreation and spiritual reflection in an idyllic environment. Their development predated all public gardens and arboreta in the United States and would help institutionalize what became a quintessential American value: that all people, even city-dwellers, deserved access to green space. Green-Wood, at the time of its founding, was believed to combine an ideal set of virtues: it was close enough for a daytrip from Manhattan and far enough away that the land around it would never get developed. Since that time, Green-Wood's surroundings have radically transformed. Now bordered by the densely populated residential neighborhoods of Windsor Terrace, Park Slope, Sunset Park, and Kensington, Green-Wood's perimeter is directly flanked not by countryside but by the lessthan-bucolic Metropolitan Transit Authority's Jackie Gleason Bus Terminal and a Con Edison substation. Over the past two centuries, the rapid development and urbanization of GreenWood's immediate surroundings have increased the cemetery's relative socio-ecological value. Sunset Park, for instance, has the least amount of green space per capita among New York City neighborhoods. In the face of urbanization and increased disturbances from climate change, Green-Wood has returned to its roots as a community-focused public garden. This is a timely and crucial return for Green-Wood's resilience as a greenspace and for supporting the ecological health of the New York City region. In recent years, Green-Wood's collection of trees and shrubs has gained increased recognition and accolades. Among other things, GreenWood has collaborated with United States Forest Service on a project that led to the discovery of a new, as yet unpublished, species of woodboring beetle (Agrilus sp.), and Green-Wood's oak (Quercus) collection is now a Nationally Accredited Plant Collection. These efforts have highlighted the institution's importance within New York City's urban forest and have supported research vital for forest preservation. But what of Green-Wood's grasslands? Literally overshadowed by the larger, more charismatic trees and shrubs, Green-Wood's expansive grasslands are by far its most complex, dominant, and resource-dependent vegetation. Alterations to the management practices of these grasslands, therefore, may stand to have the most impact on the sustainability and resilience of Green-Wood in the face of climate change. Urban Grassland Ecosystem Services Over the last twenty years, with the advent of improved data capture and analytic technologies, researchers and policy makers have become increasingly interested in quantifying the relationship between humans and the natural world. The United Nations codified the conceptual framework of this relationship, known as ecosystem services, in the Millennium Ecosystems Assessment report published at the turn of the twenty-first century. The anthropocentric view of ecosystem services asserts that the natural world serves the needs of humans in measurable ways: by regulating climate and ecosystem health, producing raw materials, supporting natural systems through chemical processes, and providing cultural benefits. Humans, however, often impact the natural world in ways that demonstrably undermine those services. Urban grasslands showcase this tension. Grasslands provide a permeable surface for stormwater to penetrate, helping mitigate runoff from increasingly frequent extreme precipitation events. They provide habitat for wildlife and space for human recreation on a cushioning vegetated surface. Concurrently, at the soil surface, a seething foundry of microbial activity sequesters greenhouse gases, fixes nitrogen, and processes pollutants. Yet, by definition, urban grasslands, especially cemeteries, are associated with regular surface disruption from mowing and excavation, which destabilizes the soil surface, increasing erosion and releasing stored greenhouse gases. Turf is considered the most widespread irrigated crop in the United States, and its management also requires fossil fuels and a multitude of chemicals, including fertilizers, herbicides, fungicides, and growth regulators. The overall maintenance cost to the American consumer is steep: according to the market research group Facing page: Urban grasslands at the Green-Wood Cemetery, in Brooklyn, provide essential ecosystem services for the surrounding high-density neighborhoods. PHOTO BY MILES ABRAMS, REDWING DRONES FRANK ROSSI 28 Arnoldia 77\/3 ? February 2020 Bermudagrass (Cynodon dactylon) spreads with aggressive rhizomes and above-ground stolons, presenting a serious management problem for Green-Wood staff. IBISWorld, households spent around $30 billion on landscape maintenance in 2019, with most of those services centered around lawncare. Expectations for high-intensity maintenance are especially pronounced at cemeteries, because they are publicly accessible landscapes of great emotional resonance, segmented by private ownership. Although ideal for a rural cemetery, GreenWood's glacially influenced topography is ill-suited for frequent mechanical mowing: the uneven ground is susceptible to scalping by mower blades and is further scraped by machinery navigating its steep slopes. The high-frequency mowing program causes surface disruption that leaves areas of bare soil and renders Green-Wood's grassland vulnerable to invasive organisms. Bermudagrass (Cynodon dactylon) is among the most aggressive invaders. Concerns about unsustainable mowing practices and the rapid expansion of Bermuda grass ultimately led to a collaboration between Green-Wood and the College of Agricultural Life Sciences at Cornell University. The threeyear partnership officially commenced in 2017 and has focused on developing intelligent and climate-sensitive strategies for grassland preservation and restoration. Intelligent Grassland Restoration Bermudagrass is a warm-season species found in the humid transition zone in the southern United States. It was likely introduced from eastern Africa through ship ballast and intentional planting as a pasture grass. With the reduced frequency of lethal winter temperatures, grasses and forbs more characteristic of warmer areas are now persisting farther north. The observed northern expansion of Bermuda grass has also been accelerated in urban environments by the heat island effect. While it is unknown how Bermudagrass arrived at Green-Wood specifically, the population has flourished over the past decade and continues to increase. Bermudagrass spreads with aggressive rhizomes and above-ground stolons, producing an impenetrable monoculture that quickly covers newer, prostrate gravestones. The success of its colonization is in part due to Green-Wood's function as an active Urban Grasslands 29 FRANK ROSSI urban grassland managers. The researchers have also deployed microclimate sensors in three areas of the cemetery that present unique vegetative characteristics due to topography and light intensity. Together, these technologies will allow the team to differentiate plant populations, measure the level of soil disturbance, and define microclimates and soil types across Green-Wood's landscape. Intelligent grassland management also relies on tools from a larger toolbox, including refined methods of soil handling, weed seedbank management, and adaptive seed mixtures. The team intiated trials in 2017, which have already yielded positive results. These last two years were the wettest in recorded history, which caused the team to assess fungal disease susceptibility among varieties in the first new seed mixture. But additional mixtures, some containing native species, have established nicely within three months and are now persisting under regular disturbance with little weed competition. These findings suggest that site-specific plant selection can help to address the persistent disturbance associated with the urban environment. FRANK ROSSI cemetery with more than one thousand burials each year. Cemetery staff excavate and relocate soil whenever a grave is dug, and Bermudagrass moves with soil and spreads into recently disturbed plots, outcompeting other vegetation. The aggressive nature of Bermudagrass creates two problems: its rapid growth requires more frequent mowing to sustain an aesthetic expected by Green-Wood's lot owners and other visitors, and the dormant stage of straw-brown vegetation during the cooler months creates a poor visual aesthetic that is highly unfavorable to the majority of cemetery stakeholders. In order to develop an intelligent grassland management system that is capable of controlling Bermudagrass at Green-Wood, the team knew that it would be essential to assess shifts in plant populations in response to maintenance. The researchers from Cornell are currently investigating the use of new agricultural technologies that can analyze satellite imagery to establish baseline Bermudagrass population levels. This technology will require on-the-ground observations to test its accuracy and will ultimately be integrated into existing mapping systems that are widely used by To better understand grassland habitats at Green-Wood, a team of researchers from Cornell and Green-Wood have installed microclimate sensors. They also conducted soil samples with colleagues from the United States Department of Agriculture's Natural Resource Conservation Service. 30 Arnoldia 77\/3 ? February 2020 The goal is not to eliminate the presence of Bermudagrass but to find ways to realistically manage its presence, a balance which would occur in concert with restoration of the grasslands. Future seed mixtures will be designed to create ephemeral flowering regimes that support specialized pollinator species and will include grass species that thrive on reduced mowing, allowing the turf to store carbon deeper in the soil profile. Colleagues at Oklahoma State University are also conducting genetic fingerprinting of Green-Wood's existing Bermudagrass population, in order to identify its unique traits. As part of a progressive adaptation strategy, we hope to establish Bermudgrass cultivars that would better meet the needs of urban grasslands in the future. Climate-Smart Mowing While distinctly rural in design, Green-Wood's landscape also reflects qualities of the lawn cemeteries that came after the rural cemetery movement: flatter areas of turf bordered by trees. Lawn cemeteries prioritized turf for its assumed ease of maintenance and its neat, uniform appearance. To maintain a manicured aesthetic, however, these lawns demand either vegetation that grows slowly and moves by underground rhizomes, or a consistent, low height-of-cut mowing regimen (often a complete rotation through the landscape every seven days). At Green-Wood, depending on the rate of growth, the maintenance of this aesthetic can equal between thirty-two and thirty-five annual mows, which require over ten thousand gallons of gasoline to complete, emitting roughly two hundred thousand pounds of carbon dioxide into the atmosphere. Operational challenges that come with this level of turf maintenance are compounded by the complexity of GreenWood's landscape, leading to worn-out vegetation and rutted soils. The team of researchers, along with GreenWood staff and contracted specialists from the landscape management company BrightView, are implementing a data-driven process that is more sensitive to climate change. They initiated a study in the summer of 2019 to reduce mowing frequency by 85 percent on approx- imately two hundred acres, chosen for their topographical features and known levels of visitation. The team tracked equipment usage to assess actual mowing times and collected detailed observation of grassland response in terms of species richness. After three months, the acreage under the experimental reducedmowing program was scaled back for several reasons: the disparate areas made it difficult to manage revised specifications; aesthetic concerns were voiced by staff members; and critical feedback came from cemetery stakeholders. While the general public voiced strong enthusiasm for the program, some lot owners saw the longer grass as a symbol of neglect. We learned that it is often best to implement changes like this gradually, allowing for increased community engagement throughout the process. Nevertheless, much was gained during this first effort. The data gathered have enabled us to align mowing frequency with growth rate, thereby permitting a slight increase in cutting height, while respecting the expectations of stakeholders. These efforts will be strengthened as we continue to sow new seed mixtures in high-visitation areas, incorporating plants that require less mowing, while simultaneously increasing species richness. Quietly Planning to Raise Awareness The association between humans and grasslands is intimate and well-established: grasslands regenerate the soil for crops, sustain grazing animals, and fulfill an innate human desire to connect with the natural world. While a growing body of research is devoted to ecosystem services provided by urban grasslands, the people who most directly interact with these urban spaces are often unaware that the landscapes are, in fact, grasslands. This is more than semantics. Seeing cemeteries, public gardens, parks, sports fields, and golf courses as urban grasslands forces a paradigm shift. In this light, the grassland is not exclusively a feature of ruralness but rather one that is present in urban environments, within walking distance. This shift can help the public see that green vegetation is everywhere: in parks and gardens, on playing fields, and in cemeteries. The urban In 2019, meadows were allowed to develop at Green-Wood. The meadows provided improved ecosystem services and lead to overall emission reductions. Photo by Art Presson. grassland concept unites all these patches of green and makes them part of a dynamic landscape. It encourages curiosity from a soccer player about the care and health of a ground on which she runs, and it transforms an apartment building's backyard lawn into an opportunity to combat climate change, simply by doing less. Raising awareness of the importance of urban grasslands like Green-Wood is a critical step toward sustainable, intelligent management. To this end, Green-Wood and Cornell are forming an Urban Grasslands Institute at Green-Wood, intended to share our findings and communicate the value of these often overlooked grasslands to a diverse urban population. The success of such an initiative is dependent on broader collaborations. The leaders of the project have invited a national group of experts from the field to consult on issues such as species selection and soil type, and will continue to look for opportunities to grow this knowledge base and expand its reach. As socio-cultural issues evolve, the Urban Grasslands Institute at Green-Wood will share information about a combination of technologies and smarter management practices, helping homeowners and grassland managers prepare for the challenges of a rapidly changing urban climate. While not explicitly articulated, one of the underlying goals of the project is to reform the discordant, one-size-fits-all model of landscape maintenance promoted by the lawncare industry, which has directly shaped the public's views of acceptable turf management. We hope to present a more nuanced model that optimizes the relationship between the biological and cultural functions of a landscape. Joseph Charap is director of horticulture and curator at Green-Wood. Sara Evans is manager of horticulture operations and projects at Green-Wood. Frank S. Rossi is associate professor of horticulture at College of Agriculture and Life Sciences, Cornell University. "},{"has_event_date":0,"type":"arnoldia","title":"Each Year in the Forest: Winter","article_sequence":5,"start_page":32,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25677","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070856b.jpg","volume":77,"issue_number":3,"year":2020,"series":null,"season":null,"authors":"Davis, Rachel D.; Hipp, Andrew L.","article_content":"Each Year in the Forest: Winter Andrew L. Hipp Illustrated by Rachel D. Davis ... Except for the point, the still point, There would be no dance, and there is only the dance. --T.S. Eliot, \"Burnt Norton\" T I he forest year has neither beginning nor end. It has, however, landmarks along its coiling journey. By December, the worms are slow in the soil. Pillbugs and woolly bears become still as the puddles freeze. Painted ladies and mourning cloak butterflies are tucked under sheaves of tree bark. At our home in the Chicago region, snow comes and goes. One weekend, we have enough to cross-country ski. By the next it has melted away. Some years, we have yet to plant the garlic. In a warm December, a few spring wildflowers start making headway on the next year. Colonies of bullet-shaped mayapple shoots emerge from the soil, leaves folded inside like tiny hands in prayer. Spring beauties sprawl beneath the leaf litter at the base of a sugar maple tree or under a rotting log, strap-like leaves curled and vulnerable, stems fragile. Foliage of the HIPP, A.L. AND DAVIS, R.D. 2020. EACH YEAR IN THE FOREST: WINTER. ARNOLDIA, 77(3): 32?39 Forest: Winter 33 false rue anemone looks as ready for the new year as it will in March. Do these individuals survive under the snow all winter long? Are they making a calculated move that will give them an edge in the spring rush? Or are they making a tactical error? No matter what year, winter arrives with unexpected greens. Dark, leathery leaves of white bear sedge grow as broad as banana peels, while the narrow-leaved sedges cluster like mop heads in the forest understory and pool in shady depressions. Fronds of the spinulose wood fern recline against the oak leaves. Seductive entodon moss carpets the decomposing boles of fallen red oaks that started growing in the mid-1800s. The moss works intermittently through winter when there are few other plants to see, collaborating across the seasons with fungi and slime molds, algae and bacteria, and mice and invertebrates to digest and break apart the fallen tree. Evergreen leaves are gearing up to spend winter under the snow, ready to photosynthesize whenever the light is bright enough to fire up their chloroplasts. They are scaling back their hours to part-time. II The soil freezes and thaws repeatedly. Under the sugar maples, bundles of needle ice form at the surface of exposed soils, each an inch or so long and packed together like fists full of glass straws. Without an insulating layer of leaves, the ice heaves knots of soil into spires reminiscent of the stone formations in southern Utah's Canyonlands. The ice melts slowly in my ungloved hand, perfectly clear near the top and middle, swimming with soil particles near the base. If the soil beneath is warm enough, the frozen clods can be brushed loose like granola off a countertop, revealing a cool, moist bed of fine crumbles and worm castings. The year pivots on the week flanking the winter solstice. We awaken to darkness and return home in darkness. On my bike ride into work, a whitefooted mouse darts across the road and disappears into a shrub. It navigates the tangle of branches, and the light I turn on it plays on its back as though Pillbug and Earthworm Deer Mouse 34 Arnoldia 77\/3 ? February 2020 White-Breasted Nuthatch Gray Squirrel Downy Woodpecker the mouse were a convict scaling a prison wall. Owls call in the morning while I walk in through the woods. In the darkness, the trees are silhouetted against the cloudy sky. The sugar maples are magnificent, messy-headed beasts with trunks as wide as picture windows that heave out of the soil and head straight up for several stories before bursting into crown. Bur oak branches stretch out even under the canopy, vestiges of an ancient savanna. Oaks and beeches hang onto a good portion of their leaves. White and red oaks are everywhere, with hop hornbeam, musclewood, black cherry, and hackberry filling in where they can. I'm freed to forget, for the space of the walk, everything I need a hand lens to see. The days hang still. There is the Christmas bird count with its coots and mallards, juncos and chickadees. The woods are filled with nuthatches, woodpeckers, golden-crowned kinglets, brown creepers, screech owls, barred owls and, down from the north, a saw-whet owl. There are always a few bluebirds. We pad around the house for the first few lengthening days of late December and January. The cardinals start to sing. Forest: Winter 35 Then snow falls and blankets everything. In the midst of a snowstorm that lasts for hours, geese may be heard calling to each other from a nearby pond. As the sun goes down, the clouds shed enormous, cottony flakes. The snow goes on all night. We awaken to a clear sky, with Jupiter swinging up above Venus's left shoulder, the moon high in the southwest, a few steps from the planets. Paper birch fruits skid across the surface of the snow. Norway spruce needles from nearby backyards pepper the drifts. If it is cold and dry enough, the wind whips the fallen snow into sharp ridges that run along the margins of fallen tree trunks, forming slot canyons that reveal duff spilling out from beneath the trunks. Snowflakes link arms, cantilevering from the tree branches. Hoar frost sprouts from the ice along the creek like moss sculpted in porcelain. Water bubbles along beneath. III Snow hides, then it records and reveals. Mammal tracks run everywhere, except during the bitterest cold. White-footed mice gallop, tails licking the surface, forepaws and hind paws paired. Their tunnels weave through the snow, leading to crystal-edged holes. A scrabbling near one end of a mouse trail captures the frustrations of a fox. Meadow voles scurry, paws alternating. But we often don't see them. They begin their paths as tunnels between grass nests and the bases of tree trunks, but they often pop up for a quick view before they dive back down. When temperatures rise, these portals through the surface of the snowpack sweat a frosty collar, and the roofs of the tunnels become thin, translucent, graying over the darkness inside. When the roofs cave in, the tunnels are etched into the melting snow. They freeze again. Then the snow melts away and is gone altogether for a few weeks. Channels appear chewed into the grass. The snow returns. This coming and going of snow is common throughout the winter in southern Wisconsin and northern Illinois, where I have lived almost all of my life, and it's fundamentally different from the persistent snowpack of the north, which insulates the ground through the coldest weeks.1 During January 2019, when temperatures hit -30?F (-34?C)--so cold that a cup of boiling Red Oak Buds 36 Arnoldia 77\/3 ? February 2020 water, tossed in the air, would vaporize into a cloud of fog before it hit the ground--the mice and voles carried on under the snow, feeding on tubers and plant stems, girdling willows. At the bottom of the snowpack, the bottom layers sublimate away, leaving a crystalline rooftop with an air gap for the mice to occupy.2 They are out of the line of sight of hawks and coyotes, though within earshot. The unlucky mouse may meet its maker crushed in the talons of a great horned owl that plucked it from the snow. Aside from that risk, the snow is the mouse's blanket and the earth its furnace. I fixate on tracks. I spent a week of 2019 mistaking skunk tracks for raccoon. Downers Grove is an exceptionally skunky town, so I should know better. Part of the blame goes to my preconceived notions about when skunks ought to be out. It's not warm enough for them, I thought. That's the downside of experience: it insinuates itself between you and what you're looking at. I'll give another part of the blame to the powdery snow, which was too airy to take more than a vague impression. But a week later, just a little warmer, and the snow was excellent for tracks. Opossum prints showed the rear-paw thumb as clearly as a textbook. I could have measured the claws on the gray squirrels and the lengths of the white-footed mice paws. The \"raccoon\" I had been following resolved clearly to skunk, with defined claws in lieu of the asymmetrical prints of the raccoons, which themselves showed up along a ditch that day with crystal clarity. Sharp-tipped maple seedlings and barren seed heads of wild leeks pierce the snow. Then the snow melts away, and I hardly notice them. A dusting over the top of a severed oak draws my attention to the white rot inside, throwing into relief the dark sutures between colonies of turkey-tail fungus devouring the wood. Dark, root-like networks of honey mushroom rhizomorphs become visible against the boles of fallen trees as the bark disintegrates. During the growing season, they invade roots of uninfected trees and work their way up beneath the bark, where the fungus infects the wood and causes decay. They aren't more prevalent in the winter, but snow masks so much that I notice things I would never see without it. IV Near the middle or end of February, winter starts to break. If it's very warm, bluebirds and mourning doves and chickadees will sing their hearts out as they shuttle around the neighborhoods. Red-bellied woodpeckers bark. Male redwing blackbirds return ahead of their potential partners and showboat around the ponds and ditches, singing from the spear-tips of the previous year's cattails and from the tops of the smaller trees. One morning a song sparrow returns: I hear it before I see it. It is a kind of springtime, but ephemeral, not the spring we planned for. In the exceptionally warm winter of 2017, fog descended on the region one February night, and we awoke to temperatures near 50?F (10?C), a winterresident robin singing, beating the sunrise by an hour and a half. I got to Forest: Winter 37 Skunk and Tracks the Morton Arboretum a bit before five. Fog hissed against the high-tension lines near the gate. I parked my bike and walked in through the unevenly blackened forest. The arboretum was a pioneer in burning oak woodlands to control weeds and promote understory diversity. The natural resources crew burns every year, and that February marked the beginning of one of the best burn seasons I've witnessed since I started working there in 2004. Raindrops hit the charred leaves at intervals, heavy, less resonant than they would have been on freshly downed leaves. This had been a particularly thorough burn, but even so, strips and patches of unburned leaves remained, and some logs that might have burned well went untouched while others smoldered for days. Flames were still darting out from the ragged end of a log. How must it have felt to come across spontaneous fire like this in the wild when it was still easier to carry coals from place to place than to start them from scratch? Over and over, groups of people must have rediscovered this mystery and felt grateful. Chorus frogs called from the lowlands near the interstate. I retrieved my bike, and on the ride into the herbarium, I heard the unmistakable whistling of a woodcock flying low overhead. He landed and uttered a single \"peent\" in the field between the planted buckeyes and the woods adjoining the lake sedge marsh. I waited a few minutes, but he had no more to say. This was early for the woodcock's return. Its migration north is perhaps more variable than other birds', who synchronize with the increasing day length. Songbirds moving through may live on insects that navigate the furrows in the warming tree bark. They search for groggy moths and butterflies, and berries still hanging from the trees. The woodcock, by contrast, follows the worms northward as they awake. Its prehensile bill is good for poking holes in the 38 Arnoldia 77\/3 ? February 2020 Woodcock Song Sparrow soil and snagging prey. Woodcocks are reputed to eat more than their weight in earthworms each day.3 The woodcock is not like the chorus frog, who can begin singing in the spring, stop when the weather gets cold, begin again, and then stop all summer long before its fall renaissance. The woodcock is different: when it returns, spring must be around the corner. The woodcock is committed to it. V We couldn't bear incessant spring exuberance. So, we are allowed a short break. Just as the most vivid dreams come when we are falling into sleep or stretching out of it, so the attenuation of stimuli in winter heightens our awareness. We notice praying mantis egg cases that we had missed in the fall. We pass a frozen pond and think of water milfoil and coontail and bladderwort on its floor, turions twisting toward March, common duckweed suspended below the surface of the ice or frozen into it, snapping turtles drifting noiselessly beneath the surface. We think back to the toads we saw moving through in June and wonder where they have buried themselves. The season meanders northward. One afternoon near the end of February, an enormous flock of sandhill cranes flies over. I may be inside with the dog, or in the garage with my bike flipped upside down, oiling the chain, when I hear their call from the south, like a sound that would have been familiar to the dinosaurs, though they never actually had a chance to meet. I run out to watch the cranes pass, impossibly high, sometimes concealed inside a cloud. They sound as loud as if they were in a park at the end of the block. They stream by, croaking, strings of them twisting away behind the tipmost point that glides on ahead. They catch sight of a marsh below and grow disoriented, suffer a few moments of uncertainty, continuing to drift northward like a cloud. Then they regroup, and then they are gone. Whatever I was thinking of when the cranes first started calling has mostly drained away, but not utterly, and the cranes are gone so soon that the thoughts flood back in. I wander back to what I was doing. We are not the same people we were last December. The cycles of freezing and thawing have heaved something loose. We are ready for spring. Forest: Winter 39 PLANTS REFERENCED Acer saccharum ? sugar maple Allium tricoccum ? wild leek Betula papyrifera ? paper birch Carex albursina ? white bear sedge Carex lacustris ? lake sedge Carpinus caroliniana ? musclewood Celtis occidentalis ? hackberry Ceratophyllum demersum ? coontail Claytonia virginica ? spring beauty Dryopteris carthusiana ? spinulose wood fern Enemion biternatum ? false rue anemone Entodon seductrix ? seductive entodon moss Fagus grandifolia ? American beech Lemna minor ? common duckweed Myriophyllum sp. ? water milfoil Ostrya virginiana ? hop hornbeam Picea abies ? Norway spruce Podophyllum peltatum ? Mayapple Prunus serotina ? black cherry Quercus alba ? white oak Quercus macrocarpa ? bur oak Typha ? glauca ? hybrid cattail Quercus rubra ? red oak Utricularia sp. ? bladderwort Praying Mantis Case Endnotes 1 Curtis, J.T. 1959. Environment. In The Vegetation of Wisconsin (pp. 25?48). Madison: The University of Wisconsin Press. 2Peter Marchand writes eloquently about this in his fascinating 1987 book, Life in the Cold: An Introduction to Winter Ecology. Hanover and London: University Press of New England. 3Ehrlich, P.R., Dobkin, D.S., and Wheye, D. 1988. The Birder's Handbook: The Field Guide to the Natural History of North American Birds (p.140). New York: Simon and Schuster\/Fireside. Andrew Hipp is the Senior Scientist in Plant Systematics and Herbarium Director at the Morton Arboretum in Lisle, Illinois. He conducts research on the origins and implications of plant diversity, with a focus on oaks, sedges, phylogenetic ecology, and trait evolution. You can read about his research at https:\/\/systematics.mortonarb.org\/lab and follow his natural history blog at https:\/\/botanistsfieldnotes.com. Rachel Davis is an independent visual artist in the Chicago area. She works at the interface of natural science, abstract painting, printmaking, and textiles, integrating the formal and empirical elements of the natural world in her work. You can see more of her work at https:\/\/artbumble.com and follow her on Instagram: @art_bumble. "},{"has_event_date":0,"type":"arnoldia","title":"Promise of Bark: Eucommia ulmoides","article_sequence":6,"start_page":1,"end_page":null,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25678","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070896f.jpg","volume":77,"issue_number":3,"year":2020,"series":null,"season":null,"authors":"Richardson, Kathryn","article_content":"RICHARDSON, K. 2020. PROMISE OF BARK: EUCOMMIA ULMOIDES. ARNOLDIA, 77(3): 40 Promise of Bark: Eucommia ulmoides Kathryn Richardson On July 1, 1910, Ernest Henry Wilson was traveling through Yunyang County, Chinathen part of eastern Sichuan Provincecollecting plant material for the Arnold Arboretum. The region is extremely mountainous, with footpaths snaking along vertiginous river valleys, through naturally formed rock tunnels, and past old fort barrack sites. Wilson photographed many large trees in the region, and his passage was crisscrossed by men carrying loads of salt and other commercial products. He photographed one of these men shouldering two large bundles of bark that were suspended from either end of a wooden rod. This was a shipment of du-zhong, a medicinal bark from the hardy rubber tree (Eucommia ulmoides), which was prescribed then, as it still is today, for kidney and liver ailments, among other health issues. Wilson never observed wild populations of Eucommia ulmoidesthe only species in its family, Eucommiaceaealthough he frequently saw two or three medium-sized trees planted near houses. Overharvesting and deforestation were likely (and continue to be) the cause for the rarity of sightings in the wild, but bark for medicine was abundant in cultivation. This medicinal use, however, was not the sole interest of botanists in Europe and North America. When the bark is harvested, dried, and gently broken, a latex-like product becomes visible. This characteristic aroused commercial interests. In 1911, Charles Sprague Sargent, the founding director of the Arnold Arboretum, wrote, in the Bulletin of Poplar Information, about considerable excitement that had arisen around the species. This is a hardy tree to which a good deal of space has recently been given in the daily papers as the 'Hardy Rubber-tree,' and as a possible source of rubber in cold climates, Sargent wrote. If true, this use would provide a considerable breakthrough, given that commercial rubber was produced from a Brazilian species, Hevea brasiliensis, which could only be cultivated in the tropics. Five years later, however, Sargent returned to the subject with a more dismal assessment, noting that the plant has been of more interest to the energetic newspaper report than it can ever be to the manufacturer of rubber goods. The Arboretum's oldest specimen of Eucommia ulmoides (accession 14538*A) grows along Linden Path, not far from the Hunnewell Building. The plant was received from James Veitch & Sons nursery in 1907 and was likely collected on an earlier Wilson expedition. The tree now graces visitors with its thick, sturdy branches that extend upward in a stair-like fashion. The bark is deeply ridged and furrowed, and has become a home for various moss and lichen. The elm-shaped leaves emerge in the spring. When gently pulled apart, the leaves reveal strings of latex within, each as thin as spiders' silk. Although Sargent's predictions about the commercial use of this latex product would prove accurate in North America and Europe, the Flora of China indicates that the rubber-like product has been successfully used for insulating electrical cables, sealing pipes, and even filling teeth. Medical research has increasingly pointed to the benefits of the bark for lowering blood pressure. Today, wild populations of Eucommia ulmoides are heavily protected. The International Union for Conservation of Nature lists the species as vulnerable to extinction in the wild and estimates that fewer than one thousand mature individuals remain in widely scattered populations, mostly on steep slopes that are difficult to access. Collectors on recent Arboretum expeditions have never witnessed the species in the wild, although, like Wilson, they have observed the trees in cultivation. Whether wild or cultivated, Eucommia provides a direct reminder about the importance of plants beyond the garden walls: as medicine, as dreams of rubber, and as livelihoods for those who harvest and share what the plants have provided. Kathryn Richardson is a curatorial assistant at the Arnold Arboretum."},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23468","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14e8528.jpg","title":"2020-77-3","volume":77,"issue_number":3,"year":2020,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Marian Roby Case: Cultivating Boys into Men","article_sequence":1,"start_page":2,"end_page":9,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25670","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070b36d.jpg","volume":77,"issue_number":2,"year":2019,"series":null,"season":null,"authors":"Pearson, Lisa","article_content":"PEARSON, L. 2019. MARIAN ROBY CASE: CULTIVATING BOYS INTO MEN. ARNOLDIA, 77(2): 2?9 Marian Roby Case: Cultivating Boys into Men Lisa Pearson Only Hillcrest farm boys, we'll soon by Hillcrest men, True and trusted citizens, we'll work for Weston then. Bound to make her greater far than she has ever been. We are the farm boys of Weston. A -- FIRST VERSE OF \"HILLCREST SONG\" t the turn of the twentieth century, Weston, Massachusetts, was a farming town that had become a country retreat for the well-to-do of Boston. With a commuter train connecting it to downtown Boston, less than twenty miles away, wealthy families had moved westward, searching for fresh air and rural activities. Among these estates arose an unconventional operation: an experimental farm, launched in 1910, by Marian Roby Case. For more than three decades, Case conducted the operations of a remarkable educational and horticultural enterprise called Hillcrest Gardens, which made a lasting impact on the boys who participated. In 1920, a Whitman Times article by Louis Graton described Hillcrest as \"a truly philanthropic institution ... where boys, any boys, may receive, under expert tutoring, up-to-date instruction in fruit and vegetable growing. These boys are also taught the rudiments of good business. They are sent out with the truck, well loaded with the choice products their own hands have helped to raise ... to sell and thus learn self-reliance.\" Marian Roby Case was born in Boston in 1864, the fourth and youngest daughter of merchant and banking executive James Brown Case and his wife Laura Lucretia Williams Case. In her youth and young adulthood, Marian and her family divided their time between their home on Beacon Street, in Boston, and their country place, Rocklawn, in Weston. After her father's death in 1907, Marian, her sister Louisa, and their mother came to live year-round in Weston. Marian had inherited about ten acres of property from her father between Wellesley and Ash Streets and proceeded to purchase other nearby plots as they became available, assembling about seventy acres of orchards and arable land over the next few years. A Passion for Horticulture It would seem that Marian Case had always wanted to farm, as apparently had her father. When Hillcrest was established, she initiated an annual pamphlet, known colloquially as the \"green books,\" given the color of their covers. In the green book for 1918, she remarked, \"[I] had inherited my father's love for the care and cultivation of land. How often in travelling have I seen my father wax enthusiastic over the well-tilled acres we have passed.\" At their Weston home, James Case had indulged in his avocation, at least during the family's summer sojourns there, by raising prize livestock for exhibition at regional agricultural fairs. The family, whose wealth came from the dry goods business, and later banking, focused their philanthropy on organizations geared towards the improvement of society. Louisa Case was a donor to the North Bennet Street School, a training program in the manual arts located in what was then a section of Boston heavily populated with recent Italian immigrants. Marian Case was an active supporter of the Hampton Normal and Agricultural Institute, of Hampton, Virginia, through its Boston association. The institute, whose most famous graduate was Booker T. Washington, sought to educate black students to create future leaders in edu- For more than thirty years, Marian Roby Case operated an experimental horticulture training program at Hillcrest Gardens, her farm in Weston, Massachusetts. ALL IMAGES FROM THE ARNOLD ARBORETUM ARCHIVES 4 Arnoldia 77\/2 ? November 2019 cation, farming, and business, and it is now known as Hampton University. Its programs stressed not only instruction in practical skills but had a deep grounding in ethical and cultural improvement. These training programs gained traction in Boston during a nationwide boom of secular and religious progressive activism in the second half of the nineteenth century, which aimed to address, among other things, rising income inequality. Andrew Carnegie famously outlined a vision for philanthropy in an 1889 North American Review article, in which he condemned ostentatious uses of wealth and urged that charitable giving should provide training and educational opportunities for the poor. James Case, for his part, attended monthly dinners hosted by the Unitarian Club, in Boston, where speakers often encouraged the affluent attendees to use their wealth for abolishing social hierarchies. In 1909, Marian Case's staff began preparing the land for the next year's farming season. The first eight Hillcrest boys were hired in 1910. This number steadily increased in subsequent years until it topped out at about twenty. The youngest were generally twelve years old, although occasionally some were younger. They worked half days for one dollar per week for the first two summers they were employed at Hillcrest. From the third summer and any summers thereafter, they worked full days and could earn up to twenty-five dollars per month. The pay was lower compared to other local farms, but each boy also received a new uniform every year, which looked rather like those of the Boy Scouts of America (an organization with complementary progressive ideals, which was also launched in 1910). The uniform consisted of two shirts, two pairs of pants, a Norfolk jacket, a tie, and a broad-brimmed hat. The boys also received a gift of educational enrichment more valuable than mere clothing in the form of lectures, study periods, journaling, report writing, and personal coaching on summer-long projects that fostered observational and writing skills. Case, following the model of the Hampton Institute, wanted to provide growth opportunities for the boys so they could develop into future leaders of their communities. We know a great deal about Hillcrest from the yearly green books, which provided a thorough review of the activities on the farm each season. The publication highlighted the reports presented by the boys during their annual convocation ceremony held on Labor Day, and these were interspersed with narratives written by Case, which provide a window into her thoughts and aspirations for her enterprise. It is interesting to see the degree to which the boys' papers became longer and more detailed as the years progressed. Some of this may be due to increased coaching that Case and her assistants were giving to the boys, but it also came from Case's desire to make the green books a resource for aspiring gardeners worldwide. We see articles by the boys to which Case had her farm manager Peter Mezitt--who founded Weston Nurseries in 1923--add additional material to explain a concept or technique more fully. Expert Instruction The boys' days were not entirely given over to farm labor at Hillcrest; Wednesday afternoon lectures were a weekly feature of the Hillcrest program. Hillcrest boy Ernest Little described them in 1935: \"One of the many advantages derived from Hillcrest Gardens is a series of instructive lectures planned by Miss Case. The program is so arranged that it includes every field of horticulture, floriculture, and botany here and abroad. They are given by leading men who are authorities in their particular line. It is with the greatest of pleasure that we welcome some of them back year after year.\" The speakers, of which there were well over one hundred by the time Hillcrest ceased operations, included a number of staff members from the Arnold Arboretum: John George Jack, who in nearly fifty years at the Arboretum was an educator, plant explorer, and dendrologist; Ernest Henry Wilson, one of the greatest plant collectors of the early twentieth century; Edgar Anderson, a geneticist and public outreach coordinator; Elmer Drew Merrill, the director (initially the supervisor) of the Arboretum from 1935 to 1946; and William Judd, a longtime propagator. Other speakers included horticultural publisher J. Horace McFarland; Arlow B. Stout, a plant breeder and research scientist Marian Roby Case 5 Over the years, the number of boys who participated at Hillcrest Gardens each summer rose to about twenty. The 1932 participants were photographed for the annual green book. at the New York Botanical Garden who spoke a number of times over the years on hybridizing and other aspects of plant propagation; John Caspar Wister, a longtime friend of Case who was a celebrated horticulturist and landscape designer; Edward Farrington, the editor of Horticulture magazine; and the Dahlia King of East Bridgewater, Massachusetts, J.K. Alexander, great-grandfather of our retired Arboretum propagator, Jack Alexander. Beginning in 1924, Case began to invite former boys back to speak at the Wednesday lectures on their experiences in business or in higher education. Brothers Joseph and E. Stanley Hobbs spoke on their respective paths into medicine and dentistry; Edmund Mezitt, whose father Peter had been employed by Case before founding Weston Nurseries, spoke about commercial horticulture; and Charles Pear lectured on his work as a weather researcher at the Blue Hills Observatory. By bringing the so-called old boys back to lecture before a new generation, Case demonstrated the success of her pedagogy at Hillcrest; boys were indeed being cultivated into active contributors to society. Cultivating Young Scholars As part of the educational component of Hillcrest, the boys were expected to keep a daily journal of their work and record their observations of the plants, insects, and weather. To this end, they were each given a notebook, pencils, and drawing paper at the start of the season. They had a daily study hour during which they could research, write about their experiences, or draw. Case worked with them personally on Fridays, critiquing their reports and coaching them on their public speaking. She also enlisted a long-serving group of local educators, including Joseph Gifford, an oratory instructor from Emerson College who worked on voice training with the boys. The summer activities culminated with the Labor Day exercises. The boys assembled and marched in carrying both the American flag and the green-and-gold flag of Hillcrest. The audience then stood for the Pledge of Allegiance, and the boys sang the Hillcrest song. Case, as mistress of ceremonies, then welcomed the guests and introduced the people who would be the judges for the boys' presentations. Each 6 Arnoldia 77\/2 ? November 2019 Chen Huanyong (left) supervised the Hillcrest boys for the first ten summers. The first enrolled participant at Hillcrest was Harold Weaver (right). of the boys read a paper they had prepared on a subject having to do with the farm. At the conclusion, the judges withdrew and chose the winners from the younger and older boys. Prizes were awarded for the papers read that day, as well as for their work in the field and in the classroom over the summer. Boys who had successfully completed one summer with distinction received a Hillcrest pin. Boys who had completed three or more summers with distinction received a pin bearing the Hillcrest motto, Semper Paratus, \"Always Ready.\" The boys' families were encouraged to attend, and in some years, the boys were allowed to invite a girl as a guest. The subjects of the boys' papers tended to repeat from year to year. There was always a report on the Wednesday lectures, the weather, and a review of the season, which would suggest that the boys chose their subjects from a list of topics provided by Case. In the 1939 green book, Case thanks Charles Sprague Sargent, the late director of the Arnold Arboretum, for his support of Hillcrest, saying, \"Soon after Hillcrest Gardens was started Professor Charles Sprague Sargent became interested in our work and helped us in many ways by giving us beautiful lilacs and other shrubs and trees, and by letting us go to him for advice.\" Sargent also persuaded Case to sponsor an essay contest for students in the Weston Public Schools. From 1921 to 1932, junior high school and high school students wrote papers on subjects suggested by Case. Unlike the summer program at Hillcrest, this essay contest was open to both male and female students, and the girls took most of the prizes over the years. Marian Roby Case 7 Cultivating Young Horticulturists From the earliest days of its operations, Hillcrest's crop production was tailored to the preferences of its customers. In the 1913 green book, Philip Coburn, who had for the previous three summers conducted door-to-door sales in Weston, writes that when the seed catalogs arrived in the winter, he and Mr. Hawkins, one of Case's full-time farm employees, chose the coming season's seeds with an eye to customer favorites. During Hillcrest's first decade, direct sales were conducted in Weston and the nearby towns of Auburndale and Waltham, by horsedrawn wagon and concurrently by truck. Produce was occasionally carried as far afield as the historic Faneuil Hall marketplace in downtown Boston. By the farm's second decade, a summer stand opened in Weston near the village blacksmith on the Post Road, and doorto-door sales in town and in Auburndale were discontinued. Instead, direct marketing was concentrated in Waltham, as the dense population allowed for the best return on their efforts. Farm production catered to this primarily Greek and Italian clientele, with tomatoes, peppers, eggplant, and parsley. Sales at the market continued until 1930 when it was decided to provide Hillcrest's produce to a Weston grocer who would then handle all the cash transactions and bookkeeping. Case was eager to trial new crops. She developed a relationship with David Fairchild of the Office of Seed and Plant Introduction at the United States Department of Agriculture and received from him new seed introductions for testing. Likewise, in 1910, Case hired Chen Huanyong (Woon-Yung Chun), a Chinese undergraduate from the Massachusetts Agricultural College in Amherst, to take charge of the boys. He worked at the farm for five seasons until 1919. Meanwhile, in 1915, Chen enrolled in the New York State School of Forestry at Syracuse University, and after his graduation he came to Harvard's Bussey Institution and studied with John Jack at the Arboretum. Chen returned to China in 1919 and later became a professor at Sun Yat-sen University. Over the years, he sent seeds for many varieties of Chinese vegetables, including eggplant, cabbage, watermelon, and bok choy, which were excitedly planted and proved popular. Seeds also came from Case's friends in Italy with whom she often wintered, including zucchini and small white eggplants. In the present day of housing subdivisions and strip malls, it is easy to forget just how rural Weston and its neighbors, Sudbury, Wayland, Lincoln, and Wellesley, were one hundred years ago. In the early years of her enterprise, Case fretted as to whether Hillcrest was cutting into the business of other local farms. She had to strike a balance between selling their produce inexpensively but not selling it at such a low price as to undercut the other farms in town. In the 1918 green book, she wrote about discussing these issues with members of the local agricultural society: \"Last spring we made thorough inquiries as to whether Hillcrest was harming the other farmers of the town and were told decidedly no. One of our well known townsmen said, `Hillcrest is doing good work. It is interfering with nobody. Go ahead.'\" The Hillcrest Boys Initially Case limited the Hillcrest program to boys from Weston but very soon expanded it to include boys from Waltham and further afield. For the period, she was remarkably progressive in her acceptance of boys for the program, welcoming sons of old Yankee families, as well as recent immigrants from southern Italy and the eastern Mediterranean, and the son of her African American butler, George Weaver. Her mentorship, respect for, and longstanding relationship with Chen Huanyong also point to her progressive ideals. In an age when children were to be seen but not heard, Hillcrest boys were encouraged to speak and make their opinions known. In fact, as Case said in 1922, \"There is no sectarian or political influence exerted at Hillcrest Gardens; each boy has a right to his opinion, whether we agree with him or not.\" Harold Weaver, the first boy enrolled at Hillcrest, participated for six seasons. He was also the first of the Hillcrest boys to go to France, in 1918, with the American Expeditionary Forces of World War I, as part of the 369th Infantry Regiment, the all-black unit nicknamed the Harlem Hellfighters. Case published an excerpt Case was photographed by Vincenzo Ruocco, in Naples, Italy, in 1929. from a letter he wrote from \"somewhere in France,\" in which he said, \"You do me honor, Miss Case when you tell me that I am the first Hillcrest Boy to come to France. My Hillcrest pin is ever with me on the lapel of my blouse. I often look at it and think how I should dislike to lose it in No Man's Land and how I hope to bring it safely from No Man's Land to Weston again so that you yourself may see the pin that has travelled 4,000 miles.\" Case went on to name two other students who were serving in the military: one as an aviator in Texas, the other in the Marines. Weaver was commissioned a second lieutenant in France, perhaps due in part to the leadership skills he learned at Hillcrest. He and the other Hillcrest boys in uniform all returned safely from their service in the armed forces at the close of the war. Case never married and had no children of her own, but she nevertheless became a second mother to about one hundred Hillcrest boys whom she guided firmly but lovingly. In 1923, she reflected on her satisfaction in one of the boy's papers, noting that his \"tribute is very pleasing to one who has tried to mother the boys, and who through a long life has seen mistakes which she feels that the training in sturdy independence, responsibility, individuality and co-operation which the boys have at Hillcrest Gardens may help to overcome.\" The loving regard with which Case was regarded by the Hillcrest boys is clear in their writ- Marian Roby Case 9 ings and later reminiscences. Case seemed to inspire affection wherever she went. She was photographed by a friend, Vincenzo Ruocco, in Naples, Italy, in 1929, and he inscribed the picture, \"A Miss Marian Roby Case madre americana, con devoto affetto offre rispettosamente ed eternamente memore il figlio italiano.\" The note roughly translates to, \"To Miss Marian Roby Case, American mother, offered with devoted affection, respectfully and eternally, from her Italian son.\" Transitions As Case became older, she began to cast about for a successor organization to take over Hillcrest. She considered the Massachusetts Horticultural Society, the University of Massachusetts, and other organizations, settling upon the Arnold Arboretum in 1942. The Hillcrest property was acquired by the Arboretum through bequests and donations by Case and her sister Louisa that occurred between 1942 and 1946. It was renamed the Case Estates and consisted of the family homestead, Rocklawn, and additional parcels of land and buildings acquired by Case. As part of the Arboretum, the property's main function was to provide additional nursery space for our living collections and to serve as a horticultural experimentation area. Weston's colder temperatures meant that plants that proved hardy there would definitely be hardy in the more temperate climate of Boston. In the 1950s and 1960s, experimental plantings and trial gardens were introduced to show plants, including herbaceous material, which would be appropriate for suburban home landscapes. The Case Estates buildings provided housing for staff and space for educational programs and public events. Case's will did not impose any restrictions on her bequests to the Arboretum, and she realized that, as times changed, the Arboretum might desire to sell the property, in which case she directed that the proceeds of the sale should be added to the general endowment. This outcome occurred in 2017 when the remainder of the property was sold to the town of Weston. In creating an educational work program for boys, Case sought to encourage future farmers, as well as leaders in whatever field a boy chose to pursue as his life's work. Women might well ask why she did not extend this program to girls. The answer is in the social mores of the era in which Hillcrest was conceived. In 1909, women were only starting to make their way into the public sphere, and it would have been very unusual for mixed groups of girls and boys to work together doing farm labor outside of a home situation. This was the era when school entrances had separate doors for boys and girls. Case grappled with what work was appropriate for the younger boys finding that the heavier farm labor was too much. Times, however, were changing. The progressive forces that inspired programs like Hillcrest were complemented by advocacy for women's rights, which extended beyond the right to vote. Pioneering women, especially the \"farmerettes\" of the Women's Land Army in Britain and the United States during World War I, led the change, and Case knew that a female horticulturist could be the match of any man. \"Sometimes I feel as if I would like to have a woman take care of my flower garden at Hillcrest,\" she wrote in Horticulture magazine in 1920. \"For as a rule women are better nurses. Men are good for spring and autumn work. They can plant, do good landscape work, go in for effects. But when it comes to the care that plants need in summer, the watching, nursing and babying I believe that women will prove better.\" She went on: \"There are some men who have this woman element to a large degree--not the feminine, there is a difference--of them, poets, artists and good gardeners are made. The marking of crosses on a piece of paper is not going to make any difference in the spirit of womanhood in either man or woman and there is still truth in the old saying that the hand that rocks the cradle is the hand that rules the world.\" Lisa Pearson is head of library and archives at the Arnold Arboretum. Her book Arnold Arboretum was published as part of the Images of America series in 2016. She had the pleasure of meeting two of the \"old boys\" from Hillcrest Gardens, Jack and Tom Williams, about fifteen years ago. They both spoke of Case in affectionate terms. "},{"has_event_date":0,"type":"arnoldia","title":"The <i>Viburnum Lentago</i> Clade: A Continental Radiation","article_sequence":2,"start_page":10,"end_page":19,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25672","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070b76b.jpg","volume":77,"issue_number":2,"year":2019,"series":null,"season":null,"authors":"Spriggs, Elizabeth","article_content":"The Viburnum Lentago Clade: A Continental Radiation Elizabeth Spriggs I arrived at the Missouri Botanical Garden's herbarium in the middle of a summer afternoon in 2014. I had spent the past two days driving and looking for Viburnum in the farm country of northern Illinois where occasional natural populations can be found in wooded roadside ditches. I was a second-year graduate student at Yale University, and it was my second collecting trip. This herbarium was an important destination because it is the second largest in the United States, and it has excellent collections of North American plants. I was planning to photograph all the viburnum specimens. I especially wanted to learn how to identify the seven viburnum species that taxonomists have traditionally grouped in the Lentago clade. A curator met me and took me to the viburnum section, where he showed me five full floor-to-ceiling cabinets. As I began to sort through the specimens that afternoon, I was totally overwhelmed: there were far too many to photograph, and I couldn't tell the species apart. The Viburnum Lentago clade is a small lineage of wide-ranging shrubs and small trees that are common in woodland edges of North America. Two members of this clade, nannyberry (V. lentago) and witherod viburnum (V. cassinoides), occur in New England and the upper Midwest. The lineage also includes the blackhaw (V. prunifolium), rusty blackhaw (V. rufidulum), and possumhaw viburnum (V. nudum), which have more southern distributions. Walter's viburnum (V. obovatum) occurs primarily in Florida, and V. elatum is restricted to Mexico. The species in this lineage all flower in the early spring and produce large blue or black bird-dispersed fruits. SPRIGGS, E. 2019. THE VIBURNUM LENTAGO CLADE: A CONTINENTAL RADIATION. ARNOLDIA, 77(2): 10?19 V. rufidulum V. prunifolium V. obovatum V. elatum core Lentago clade V. lentago V. cassinoides V. nitidum V. nudum complex V. nudum One of the most notable features of the group is that its members look very similar to one another, and nearly all the traits that distinguish the species are subtle. After my first day at the herbarium, I reread the two best treatments of the Lentago clade: Thomas Jones's 1983 graduate thesis and Waldo McAtee's A Review of the Nearctic Viburnum. Both taxonomic keys are full of equivocal phrases like \"leaves usually short-pointed or rounded apically,\" \"petioles more or less crinkly-margined,\" and \"veins less scurfy, usually glabrous.\" Resigned to the fact that I might not be able to tell the species apart, I decided to photograph as many specimens as I could with the hope that I would be able to figure them out once I was back in New Haven. Over the next couple of years, I spent many weeks collecting viburnums from wild populations in the eastern United States, collecting 330 individuals. I spent many more weeks looking at herbarium specimens. I did eventually learn to tell the species of the Lentago clade apart. Many botanists know the feeling: familiar species become easy to identify, but it is almost impossible to describe what is dis- tinctive about them. Once I knew the species, I understood the reason for ambiguous descriptions in the taxonomic keys. My own ways of identifying the species are just as difficult to describe--like picking out friends in a crowd of people. This detailed work on a small plant lineage led to surprising results and new insights into the biology of the clade. We even rediscovered a species, Viburnum nitidum, in the south eastern United States, which has been ignored for most of the past two centuries. It turned out that a lot can be learned from a group of plants that was known even to Linneaus, a lineage collected hundreds of times and studied by generations of taxonomists. History of the Clade Although most viburnums today occur in temperate or boreal environments, phylogenetic evidence suggests that the ancestors of Viburnum, and of the Lentago lineage specifically, lived in the warm temperate forests of Asia. Then, around thirty to thirty-five million years ago, Lentago moved into North America Taxonomists show the evolutionary relationships between species using phylogenetic trees. Here, a tree for Viburnum species in the Lentago clade reveals two primary subgroups: Species in the core Lentago clade, like V. prunifolium (left), produce flowers and fruit in unstalked clusters. Species allied with V. nudum (above) are borne on stalks. ALL PHOTOS BY THE AUTHOR UNLESS NOTED 12 Arnoldia 77\/2 ? November 2019 (Landis et al., in prep). The only close relatives of the Lentago clade are V. punctatum and V. lepidotulum, which both live in warm forests in Southeast Asia, many thousands of miles of away. Because no closely related species occur in Europe or Northern Asia, it is impossible to be sure how the Lentago clade's ancient migration occurred. Researchers, however, have found a fossilized viburnum pollen grain in Iceland, which has distinctive characteristics typical of species in the Lentago clade. This suggests that Lentago was likely present in Iceland about fifteen million years ago and may have migrated to North America through Europe, over the North Atlantic Land Bridge (Landis et al., in prep). Once in North America, the Lentago clade split into two lineages: the core Lentago clade and the Viburnum nudum species complex. The core Lentago includes five species: V. lentago, V. prunifolium, V. rufidulum, V. obovatum, and V. elatum. The V. nudum species complex is a small, variable lineage, which I'm calling a \"species complex\" because when I started working on it, there was a lot of uncertainty about how many species were included and how they were different from one another. Like most of Viburnum, the species in this complex bear fruit on stalked inflorescences (umbel-like compound corymbs), but in the core Lentago clade, the inflorescences are sessile or unstalked, which means that leaves are produced immediately under the inflorescence branches. Each of these two lineages has radiated into habitats that today span the range of eastern North America from central Florida to Nova Scotia. Wild Hybridization One of the most enigmatic species of the Lentago clade is the blackhaw, Viburnum prunifolium. This species occurs in a geographic area that overlaps with both V. lentago and V. rufidulum, and it is intermediate between them in many traits. Joe Brumbaugh and Arthur Guard, who observed the overlapping ranges of all three species in Indiana, concluded, in 1956, that repeated backcrossing between these species--a process known as introgression-- could be a significant cause of taxonomic confusion. Botanist Linda Rader went further and argued, in 1976, that V. prunifolium might actually be a hybrid species formed by an ancient hybridization event between V. lentago and V. rufidulum parents. Although V. lentago and V. rufidulum do not come in contact today, it is reasonable to imagine that they might have shared a geographic range in the past and might have had opportunities to hybridize. Each of these theories about hybridization is supported by the fact that hybrids among the species are possible. The cross between Viburnum lentago and V. prunifolium is known as V. ? jackii. Alfred Rehder, a taxonomist who worked at the Arnold Arboretum for much of his career, proposed the name in honor of his colleague John George Jack, who, in 1908, noticed plants in the Arboretum that appeared intermediate between V. lentago and V. prunifolium and assumed that they were spontaneous hybrids. Although those individuals are no longer at the Arboretum, a specimen of V. ? jackii growing at the Morton Arboretum was obtained from the Arnold Arboretum and is likely to be from the same original lineage. The only documented set of controlled crosses between Viburnum lentago and V. prunifolium was carried out by Donald Egolf, a graduate student at Cornell University who would become a leading research horticulturist at the United States National Arboretum. His findings, in 1956, found that a cross between V. lentago and V. prunifolium yielded twentyeight seeds and twenty-six plants, roughly the same number as crosses between only V. lentago. These numbers are surprising because they suggest that intrinsic barriers that could prevent hybridization are very low for this species pair: if an insect transported pollen from V. lentago to a V. prunifolium flower, the V. prunifolium would likely produce fertile seeds. All of this indicates that hybridization is possible and could even be common in natural populations where both species occur. In order to test for hybridization and get a better understanding of these species, I went on a series of road trips to collect leaf samples from across the range of Viburnum lentago, V. prunifolium, and V. rufidulum, including areas where multiple species occur. At each natural population, I collected leaves to use for morphological measurements and leaves to use for DNA extraction and sequencing. Across all of these areas, I found no evidence of hybrid zones or ARNOLD ARBORETUM, US FOREST SERVICE, AND GIS COMMUNITY Viburnum 13 The overlapping ranges for three species in the core Viburnum Lentago clade raise questions about how these species remain distinct. Shown with V. lentago in blue, V. prunifolium in yellow, and V. rufidulum in green. of widespread gene flow among species. While the species sometimes appear morphologically similar to one another, it turned out that this variation is not related to genetic structure or hybridization. That is, in cases where V. rufidulum looked somewhat like V. prunifolium, genetic sequencing showed that it was still 100 percent V. rufidulum. Overall, our sequencing found that V. prunifolium did not originate through hybridization. Instead, V. prunifolium is sister to the southern V. rufidulum, and the northern V. lentago is a more distant relative (Eaton et al., 2017; Spriggs et al., 2019). Out of all the 180 individuals sequenced, only two appear to be admixed (have genes from two different species). These two individuals are from a small population in northern Kentucky where Viburnum prunifolium and V. rufidulum occur together in a rocky roadside woodland, five miles east of the Kentucky River. These individuals were morphologically similar to V. rufidulum (I labeled them as V. rufidulum when I collected them), but when sequenced, they appeared to be half V. prunifolium and half V. rufidulum. Floral Timing Whenever hybridization is possible but rare, it suggests that something is acting to prevent or eliminate hybrids, in other words, mechanisms of reproductive isolation. Hybridization in natural populations has several possible outcomes. One possibility is that hybridization between two species will be so common that the species will eventually merge to become a single species. At the other extreme, if hybrids between two species are unfit, natural selection can cause species to evolve ways of avoiding one another. In some plant lineages, closely related species have different numbers of chromosomes, often leading to inviable offspring. Another common way plants avoid interbreed- 14 Arnoldia 77\/2 ? November 2019 Flowering time may explain why few natural hybrids occur between Viburnum prunifolium (left) and V. rufidulum (right). The author photographed both species on April 27, 2015, in Indiana. These temporal differences were later supported by analysis of herbarium specimens, which revealed a nine- to ten-day difference between flowering times for these closely related species. ing is with flowers that evolve to attract different pollinators. In the Lentago clade, however, the species have the same number of chromosomes (Egolf, 1956), and their flowers are extremely similar (Donoghue, 1980). How, then, do these species in the core Lentago group maintain their separation? The answer seems to lie in subtle habitat differentiation and phenological timing. Brumbaugh and Guard, in 1957, described habitat distinctions for the three overlapping species based on their work in Indiana: \"V. lentago occupies poorly drained areas; V. prunifolium, moist borders of woods, and V. rufidulum, dry rocky slopes.\" This characterization matches my own experience in the field, although I have also found that it is not difficult to find areas with two or more of the species occurring together. This kind of habitat differentiation could decrease how often the species flower in close proximity to one another. If the species are also adapted to slightly different habitats, hybridization might be disadvantageous because it could separate beneficial traits or cause beneficial traits to be lost altogether. Even more intriguing, several authors who have observed these species in the field or at arboreta sometimes mention offset flowering times. Viburnums typically flower once a year for only ten to fourteen days, and flowering time is often very synchronized within species (Donoghue, 1983). Rehder, in 1920, wrote that Viburnum prunifolium flowers about a week before V. lentago, and Rader, in 1976, noted that V. prunifolium flowered about two weeks before V. rufidulum. My doctoral advisor Michael Donoghue observed this same pattern, where V. prunifolium flowers about a week ahead of the other two species, while he was conducting his own dissertation research at the Arnold Arboretum (Donoghue, 1980). To investigate whether flowering time varies between the species in wild populations, I drove through Ohio, Indiana, and Kentucky in the early spring of 2015. In this region, V. prunifolium was very common and V. rufidulum less so, but over the course of a week, I found multiple populations of each species and consistently found V. prunifolium in full flower while V. rufidulum had green buds. Viburnum 15 Anecdotal evidence from single locations (even wild populations) is helpful but not all that convincing because each of these species occurs across a large geographic area, and each year populations experience a variety of climate conditions. To try to understand patterns of flowering time, I used herbarium specimens, which were excellent because the species are relatively common and have been collected hundreds of times from across their ranges. Even more importantly, most specimens are reproductive, meaning they have flowers or fruits, so we found plenty of suitable material, representing a large number of years. With the help of Caroline Schlutius, an undergraduate researcher, I found 1,379 flowering specimens that spanned the range of each of the species. By examining these specimens, we found clear geographic patterns in flowering time and consistent differences in flowering between species. First, we found that within each species, southern populations in warmer climates flowered earlier than northern populations in colder climates. More surprisingly, we found small but consistent differences in flowering time that remained remarkably constant across the regions where species co-occurred. In any given location, a nine- to ten-day difference in flowering time occurs between species, with a sequence that matches previous observations-- Viburnum prunifolium flowers before V. lentago and V. rufidulum. Because viburnums only flower for ten to fourteen days total, this small offset can dramatically decrease the opportunities for pollen transfer among plants. To be clear, these findings do not suggest that all of the individuals will flower in sequence every year in every location, only that in any given place for a particular year, the majority of individuals of one species will flower ahead of the majority of individuals of the other. Hidden in Plain Sight A second major focus of my work on the Lentago clade was the Viburnum nudum species complex. For the V. nudum complex, I wanted to sort out how many species there are, where they occur, and whether any traits consistently differentiate the species. In my first year collecting viburnums, I started in Florida and drove north. I was specifically targeting the V. nudum species complex, but it was very hard to find. I was surprised because viburnums are easy to find in New England, and I had expected to encounter populations driving down sideroads or in the state parks where I had permits to collect. This was not the case, and for future trips, I researched locations extensively using herbarium specimens and talking with local botanists. The first year, I collected a couple of individuals near Gainesville and then didn't find V. nudum again for a full week. The populations I had collected in Florida were in sandy soil along shallow streams, and I was looking for similar habitats as I made my way up the East Coast. I was driving on a small road along the edge of a black-water swamp in coastal South Carolina when I found the next population. A small group of viburnums was strung out along the edge of the road, several inches deep in muddy water, near a swamp with bald cypress (Taxodium distichum) and water tupelo (Nyssa aquatica). Not only was this a totally different environment than I was expecting but the plants looked different. The Gainesville plants had seemed delicate: they had small, narrow leaves, and the inflorescences bore bright pink and more mature black fruits simultaneously. This South Carolina population had thick leaves, larger than my hand, and pale green fruits. After several more collecting trips over the next several years, allowing us to sequence individuals from many populations, we discovered that these habitats are both typical for the V. nudum complex, but they contain totally separate genetic lineages, each adapted to its own environment. These results were surprising because they are at odds with the generally accepted taxonomy of Viburnum nudum species complex, which dates to the eighteenth century. Linnaeus described V. nudum in the first edition of Species Plantarum, published in 1753, and added V. cassinoides in the second edition, in 1782, distinguishing V. cassinoides by its leaf shape. Then, in 1789, William Aiton, the first director of the Royal Botanic Gardens, Kew, proposed two additional species, V. nitidum and V. laevigatum. Since then, more than eight other names have been proposed for segregates within the complex (McAtee, 1956), but none 16 Arnoldia 77\/2 ? November 2019 As the author dug into the puzzling taxonomy of the Viburnum nudum complex, fieldwork revealed that the plants occurred in three distinct habitats. Most surprisingly, those in sandy soils in Florida (above) proved to be a long overlooked species, V. nitidum. Viburnum 17 of these have been widely recognized. In recent floras, only two species or subspecies are recognized: V. cassinoides in the North and V. nudum in the South (Jones, 1983; Small, 1933; Gleason and Cronquist, 1991; Ferguson, 1966; Radford et al., 1968; Strausbaugh and Core, 1978; Weakley, 2012). When we sequenced DNA of eighty individuals, we found three different lineages in the Viburnum nudum species complex, all of which seem to be evolving independently. Most importantly, we found that the way V. nudum is typically described makes it paraphyletic, meaning that the name refers to a partial evolutionary lineage. The oldest genetic split in the V. nudum complex lineage is not between the northern and southern populations (traditionally V. cassinoides and V. nudum). Instead it is between the large-leaved populations that occur in swamps and the rest of the complex (including sandy stream populations and northern populations). Our genetic data show that each of these three lineages is distinct and evolving independently, and therefore all three deserve to be recognized as species (Spriggs et al., 2018). Linnaeus's V. nudum matches the large-leaved species that occurs in swamps; V. cassinoides corresponds to the northern species in our analyses; and the sandy-soil species seems to match Aiton's V. nitidum. After surveying herbarium specimens from across the Southeast, we determined that Viburnum nitidum is mostly restricted to the coastal plain. From Florida it extends up to North Carolina, along the coast and in the Sandhills region, then west into the eastern edge of Texas. V. nudum is more widespread and occurs throughout the coastal plain and the Piedmont, from Delaware to Arkansas. The habitats of these two species are interdigitated across the Southeast, and the species occur in close proximity to one another frequently yet remain distinct and do not hybridize. Our findings support recent arguments that the flora of the North American Coastal Plain is under-described, meaning it is more diverse than the current taxonomies suggest (Sorrie and Weakley, 2001; Noss et al., 2015). It seems that this disregard was not always the case. Over the past century, field botanists, particularly southern botanists like William Ashe, of North Carolina, or Alvan Chapman, who spent most of his career in Georgia, recognized subtle variation in the habitats of the coastal plain and described many species. These proposed species have been systematically ignored or lumped into larger widespread \"species\" that may turn out to be paraphyletic. Genetic sequencing may vindicate at least some of these descriptions, as was the case for Aiton's V. nitidum. Significance of this Clade It is reasonable to question whether it is important to know how many species are in the Lentago clade or to know how exactly they are related to one another. If Viburnum nitidum is very similar to V. nudum, does it really matter that it has a separate name? Is it useful to know whether V. prunifolium originated as a hybrid species? Does knowing the recent evolutionary history of these species have any broader implications? I believe that all this lineage-specific knowledge about species limits, occurrences, and history is important for conservation and also provides insight into the ecology and evolution of North American forests. For one thing, species identities are fundamentally important because they affect how species are recognized and valued, and whether they are conserved. Some might view the long list of proposed (and subsequently ignored) botanical names for segregates of the Viburnum nudum complex as wasted effort, but I view these names and descriptions as essential contributions, stepping-stones leading along a path to accurately characterize plant diversity. The numerous common names for V. nudum, including witherod viburnum, possumhaw, wild raisin, and Appalachian tea, suggest that even nontaxonomists knew these plants and attempted to differentiate them. With genetic sequencing, we have new opportunities to get this right, to rigorously test species limits so that the current names and species descriptions are accurate. This is especially important because relatively few plant lineages in North America (or elsewhere) have been studied with fine-scale genetic sequencing, and there is much to learn. Accurate species descriptions are also an essential starting point for ecological studies that seek to understand species distributions 18 Arnoldia 77\/2 ? November 2019 Finally, understanding the evolution of this small lineage can provide important insights into the evolution of the North American flora more generally. Each of the two major lineages of the Lentago clade diversified in eastern North America over many millions of years, and each lineage has differentiated into a series of morphologically similar but ecologically distinct species. Similar patterns of slow diversification into subtly different species are also apparent in other North American lineages like maples (Acer), dogwoods (Cornus), or ash (Fraxinus). Our work showed how the species of the Lentago clade have persisted through climate fluctuations associated with glaciation and provided evidence on one of the mechanisms (flowering time) that these species use to maintain their separation. Tensions around species identifications have hindered our understanding of the Lentago clade for over a century, but after years of observation in the field and extensive genetic sequencing, the species that had seemed incomprehensible to me at the SUZANNE MROZAK or interactions. If Viburnum nitidum and V. nudum are considered as a single entity, their distribution might be confusing or bimodal because it would include two very different habitat types. If, instead, V. nitidum is considered alone, its habitat preferences would likely be much more specific, and it might even be a useful indicator species that could serve as a quick way to identify a particular habitat or plant community. Similarly, species often differ for traits that are not readily apparent to human observers, and these can be critical in shaping interactions with insects or other species. Imagine, for instance, that there was reason to suspect that V. nitidum was a host species for a rare caterpillar, but in an assessment of this relationship, a researcher sampled V. nudum thinking it was the same as V. nitidum. This kind of mistake could lead to inconsistent or misleading results. Species are an essential unit for many ecological studies, and when species descriptions are inaccurate, there can be significant consequences. Taxonomic research reveals that all plants--even common garden denizens like Viburnum cassinoides (above)-- provide a record of millions of years of plant evolution. Viburnum 19 Missouri Botanical Garden herbarium became clear. Many other plant lineages that form the foundation of the flora of eastern North America likely have similar histories of subtle differentiation and persistence, promising countless stories waiting to be revealed. Landis, M.J., D.A.R. Eaton, W.L. Clement, B. Park, E.L. Spriggs, P.W. Sweeney, E.J. Edwards, and M.J. Donoghue. In prep. Joint estimation of geographic movements and biome shifts during the global diversification of Viburnum. References Noss, R.F., W.J. Platt, B.A. Sorrie, A.S. Weakley, D.B. Means, J. Costanza, and R.K. Peet. 2015. 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Chapel Hill, NC: The University of North Carolina Press. Clement, W.L., M. Arakaki, P.W. Sweeney, E.J. Edwards, and M.J. Donoghue. 2014. A chloroplast tree for Viburnum (Adoxaceae) and its implications for phylogenetic classification and character evolution. American Journal of Botany, 101: 1029?1049. Rehder, A. 1920. New species, varieties and combinations from the herbarium and the collections of the Arnold Arboretum. The Journal of the Arnold Arboretum, 2(2): 121?128. Donoghue, M.J. 1980. Flowering times in Viburnum. Arnoldia, 40(1): 2?22. Sorrie, B.A. and A.S. Weakley. 2001. Coastal plain vascular plant endemics: phytogeographic patterns. Castanea, 66: 50?82. Donoghue, M.J., B.G. Baldwin, J. Li, and R.C. Winkworth. 2004. Viburnum phylogeny based on chloroplast trnK intron and nuclear ribosomal ITS DNA sequences. Systematic Botany, 29: 188?198. Eaton, D.A.R., E.L. Spriggs, B. Park, and M.J. Donoghue. 2017. Misconceptions on missing data in RADseq phylogenetics with a deep-scale example from flowering plants. Systematic Biology, 66(3): 399?412. Egolf, D.R. 1962. A cytological study of the genus Viburnum. Journal of the Arnold Arboretum, 43(2): 132?172. Egolf, D.R. 1956. Cytological and interspecific hybridization studies in the genus Viburnum (Unpublished doctoral dissertation). Cornell University, Ithaca, NY. Ferguson, I.K. 1966. The genera of Caprifoliaceae in the southeastern United States. Journal of the Arnold Arboretum, 47(1): 33?59. Gleason, H.A. and A. Cronquist. 1991. Manual of vascular plants of Northeastern United States and adjacent Canada (2nd ed.). Bronx, NY: New York Botanical Garden. Jones, T.H. 1983. A revision of the genus Viburnum section Lentago (Caprifoliaceae) (Unpublished doctoral dissertation). North Carolina State University, Raleigh, NC. Small, J.K. 1933. Manual of the Southeastern Flora. New York: Author. Spriggs, E.L., C. Schlutius, D.A.R. Eaton, B. Park, P.W. Sweeney, E.J. Edwards, and M.J. Donoghue. 2019. Differences in flowering time maintain species boundaries in a continental radiation of Viburnum. American Journal of Botany, 106(6): 833?949. Spriggs, E.L., D.A.R. Eaton, P.W. Sweeney, C. Schlutius, E.J. Edwards, and M.J. Donoghue. 2018. Restriction-site-associated DNA sequencing reveals a cryptic Viburnum species on the North American Coastal Plain. Systematic Biology, 68(2): 187?203. Spriggs E.L., W.L. Clement, P.W. Sweeney, S. Madri??n, E.J. Edwards, and M.J. Donoghue. 2015. Temperate radiations and dying embers of a tropical past: the diversification of Viburnum. New Phytologist, 207(2): 340?354. Weakley, A.S. 2012. Flora of the Southern and MidAtlantic States. Retrieved from http:\/\/www. herbarium.unc.edu\/flora.htm The map in this article was created using Esri, USGS, USFS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, lntermap and the GIS user community. Elizabeth Spriggs is a Putnam Fellow at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Resolving the Enigma of Rainforest Biodiversity","article_sequence":3,"start_page":20,"end_page":29,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25671","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070b726.jpg","volume":77,"issue_number":2,"year":2019,"series":null,"season":null,"authors":"Ashton, Peter Shaw","article_content":"Resolving the Enigma of Rainforest Biodiversity Peter Ashton A berdeen, Scotland, as it was when I arrived as a professor, in 1966, at the cusp of the North Sea oil bonanza, was a very different coastal town from those in Brunei and Sarawak, where I had been based for eight of the previous ten years, while conducting fieldwork in the rainforests of Borneo. At 57? north latitude, the summer skies in Aberdeen never completely darken and native tree species are few, but the University of Aberdeen had a long tradition of research and instruction in tropical agriculture, including tea and rubber. As a new professor, I found colleagues with common interests, who shared my enthusiasms and encouraged me to continue in my research. A central puzzle had grown in my mind: How can so many species of giant sessile organisms--rainforest trees--apparently coexist in stable mixture and how did these communities originate? Elmer Merrill, celebrated botanist of East Asian floras and director of the Arnold Arboretum from 1937 to 1946, reasoned that the extraordinary species diversity of tropical and warm temperate forests in East Asia suggested an origin there for all flowering plants. Paleontological research and the growing science of molecular phylogeny would ultimately indicate the reverse--suggesting that the Aisian tropics were primarily devoid of broadleaf evergreen rainforests until the current tree families invaded in more recent evolutionary history-- yet the warm, wet equatorial climate of the tropical Far East appears, nevertheless, to be optimal for tree growth and survival. Around 4,500 plant species are known to occur in northern Borneo. ASHTON, P. 2019. RESOLVING THE ENIGMA OF RAINFOREST BIODIVERSITY. ARNOLDIA, 77(2): 20?29 Rainforest Biodiversity 21 Understanding how rainforest tree species sustain a stable mix within their communities and why these communities vary in species diversity was more than theoretical interest: If species are distributed at random within their soil-defined communities, how could consistent and predictable protocols for silvicultural management following timber harvesting be devised? At the time, research in temperate ecosystems was revealing that populations of tree species were remarkably genetically variable. This was the opposite of the widespread assumption that, for rainforest trees in species-diverse communities, self-pollination would prevail, resulting in low genetic variability within populations. As Charles Darwin foresaw, natural selection depends on the existence of diversity, which we now know to be genetic and heritable. Genetic uniformity within populations would imply lack of selection and suggest that species sharing the same habitat are ecologically complementary, surviving together through the random consequences of their seed dispersal. We decided to adopt a broad approach to explore these issues, combining crosspollination experiments with comprehensive observation of the reproductive biology of selected species, tracking them from bud formation to seed dispersal. We would bag the flowers after carefully brushing pollen onto the stigmas and then examine the genetic consequences of these crosses, comparing the genetic variability of the seedlings to the variability sampled among trees in the broader population. This research came with formidable challenges: the first was to find a safe way to ascend a 150-foot (45-meter) tree to its outer twigs in order to manipulate cross-pollination using a squirrel-hair artists brush. We were fortunate from the start. The North Sea oil was beginning to flow. I found an enterprising oilfield engineer who would work with me to devise a means of reaching the flowers through tree prosthesis: Three telescoping aluminum alloy booms, each 15 feet (4.6 meters) long, which were light enough to be carried into the forest to a tree about to flower. Combined with a cable, ropes, a simple manual dockside winch, and a boatswain's chair, the booms allowed a researcher to be lifted to the place of operation. But who among us academics, approaching middle age, would volunteer? The solution was obvious: find some students! So it was that I managed to write a persuasive grant proposal to an independent foundation created by the Unilever Corporation, from which I succeeded in gaining the necessary support. Six graduate scholarships were awarded for Malaysian students who had the necessary combination of fieldwork interest and some experience, curiosity, scientific acumen, and derring-do. Three women and three men were selected. Each student variously focused on the reproductive biology of both an emergent and a subcanopy tree species. Fieldwork was carried out at the Pasoh Forest Reserve, in central Peninsular Malaysia, about one hundred miles southeast of the capital city, Kuala Lumpur. At that time, the pollinators of dipterocarps--trees in the family Dipterocarpaceae, which dominate the overstory of these rainforests--were unknown. But in the second year, student Chan Hung Tuck collected inflorescences from a tree, sealed them in a plastic bag, and took them back to his lab at the university. To his amazement, when he opened the bag the following morning, he found several tiny insects and holes in the buds from which they had apparently escaped: thrips. Another student, Simmathiri Appanah, immediately got to work, dangling sticky plastic bottles in the canopy to trap insects. Together, their work confirmed that thrip eggs were laid in the flower buds and that thrip populations increased day by day as the dipterocarps developed buds en masse. When the open corollas fell to the ground beneath, a haze of tiny organisms was released, the insects batting their oar-like wings in the humid air. Petals are one of the very few plant organs not chemically defended in the rainforests, so they are like vegetarian McDonald's hamburgers sustaining rainforest insect diversity. The genetic work yielded unexpected results as well. The Pasoh population of the dipterocarp Shorea leprosula, by then the chosen emergent Graduate student Chan Hung Tuck uses a custom-designed boom to perform cross-pollination experiments in the canopy of Shorea leprosula. ALL PHOTOS COURTESY OF THE AUTHOR ARNOLD ARBORETUM AND GIS COMMUNITY 22 Arnoldia 77\/2 ? November 2019 INDIA Fushan Lienhuachih TAIWAN Nanjenshan CHINA BANGLADESH Dinghushan Nonggang Heishiding Xishuangbanna Hong Kong MYANMAR LAOS Doi Inthanon Palanan Hainan Huai Kha Khaeng THAILAND Mo Singto VIETNAM PHILIPPINES CAMBODIA Ngardok PALAU Bidoup Khao Chong Danum Valley MALAYSIA Lambir Pasoh E Q U A T O R Sabah BRUNEI Kuala Belalong Sarawak Bukit Timah SINGAPORE Borneo INDONESIA Ashton's research plot at the Pasoh Forest Reserve became one of sixty-seven sites in the Center for Topical Forest Science, now known as the Forest Global Earth Observatory (ForestGEO). Sites in Southeast Asia and southern East Asia are marked in green. for all our research, proved to have a genetic structure remarkably similar to the average temperate broadleaf canopy tree, with variability relatively low among neighboring trees but increasing to a population average at a radius of about 330 feet (100 meters). This is consistent with the maximum usual distance of dispersal of the winged dipterocarp fruit and the possible distance that thrips might be wafted by daily air turbulence within the sunny forest canopy. Later studies by others have confirmed that this phenomenon is close to the general rule, although some emergent dipterocarps and trees in other families attract pollinators that forage over long distances. Examples we examined included tree species visited by the giant Asian honey bee (Apis dorsata) and others visited by a cave-roosting, nectar-feeding bat (Eonycteris spelaea). Many of these same tree species bear large comestible fruit, the seeds of which are dispersed by mammals and large birds. It was becoming clear, however, that a preponderance of minute, small-winged pollinators, wafted aloft between distant conspecific individuals in the forest cornucopia, was maintaining genetic diversity across widespread habitats. Critically important, Robert MacArthur and my future Harvard colleague Edward O. Wilson had shown, in their 1967 book, The Theory of Island Biogeography, that animals on islands accumulate species at rates of immigration and extinction that vary with the area of the island: an increase of area by 90 percent is required to double the size of a fauna. Dipterocarps, however, did not follow these predictions. Whereas the large island of Sumatra includes just over 100 dipterocarp species, there are 158 species in Peninsular Malaysia, only one quarter of the area, while Borneo, one and one half the area of Sumatra, contains 270 species. The overall tree floras are consistent with these figures. As a consequence, the relationship between the number of motile organisms--animal species-- and the land masses they occupy, predicted by Rainforest Biodiversity 23 Ashton operates the boom equipment at the Pasoh Forest Reserve. MacArthur and Wilson, are rarely achieved by plants, especially trees, on account of the diversity of soils within which species are confined by interspecific competition. I joined the staff of the Arnold Arboretum in 1978, and my time as director was, for a while, fully occupied with pressing issues at Jamaica Plain. But in 1982, I attended a meeting on tropical forest ecology with my former Aberdeen student Ian Baillie, a tropical forest soil scientist. We presented a paper showing how soil nutrients governed forest community composition at topographic and geologic spatial scales in lowland Borneo. I had, by that time, become convinced that tree species in hyperdiverse rainforests were niche specific, and I published a pioneer paper to that effect. Also presenting at the conference was someone unfamiliar to me from the University of Iowa, Steven Hubbell. His paper gave me a shock! His approach and conclusion were dramatically different from my own. He had established one large tree-demography plot that covered 124 acres (50 hectares) on a relatively uniform and gentle slope on Barro Colorado Island, a research island in the Panama Canal, administered by the Smithsonian Tropical Research Institute. Steve had censused, tagged, mapped, and identified all trees larger than 0.4 inches (1 centimeter) in diameter, numbering over three hundred thousand trees--a staggering figure. From this, he had convincingly shown that the spatial distributions of the species were consistent solely with the constraints of limited seed dispersal from parent trees. He concluded that the species were ecologically complementary with one another and were therefore consistent with the geographical expectations of the theory of island biogeography after all. Following his session, I was in a state of shock. How then, if at all, could the hundreds of tree species sharing a common habitat distrib- 24 Arnoldia 77\/2 ? November 2019 Stuart Davies is now the director of the Forest Global Earth Observatory (ForestGEO). He holds the Frank Levinson Chair in Global Forest Science and is a senior staff scientist at the Smithsonian Tropical Research Institute. ute themselves independent of any variation in their physical habitat? I introduced myself and suggested we retire to a neighboring pub to hammer out our differences over a pint or two. We realized that the disagreement likely arose from our different sampling methods: my small plots were distributed across variable landscapes on a regional scale while his single large plot represented a uniform habitat. My plots were too small to detect local patterns, nor had I mapped my trees, while his plot may have been too small to detect habitat-related floristic change in relation to topography, nor had he sampled soils. We agreed that the way forward was to replicate his large plot on the other side of the world. I gained the support of my friend Salleh Mohd Nor, the director of the Malay- sian Forest Research Institute, to establish a 124-acre (50 hectare) plot on the gentle topography of Pasoh Forest Reserve. Then, Steve and I successfully persuaded the National Science Foundation to fund it. We would use identical census protocols at both sites and recensus each every five years. The aim was to resolve the central, as yet not fully resolved question: To what extent are rainforest tree species niche specific and to what extent are they spatially restricted by their limited seed dispersal? I was soon in luck again: the United States ambassador in Thailand, John Gunther Dean, was a resource economist, and he recommended that the State Department should host a regional conference in East Asia on research priorities for the sustainable development of Rainforest Biodiversity 25 natural resources. The State Department was looking for someone to orchestrate it. I jumped at the opportunity. With support from the National Science Foundation and Agency of International Development, I then toured the region seeking advice from friends and colleagues. It was not difficult to gain consensus for the concept of a regional network of representative forest community samples. The sites would follow Steve's protocol, varying in area such that each captured at least one hundred individuals of half the species represented. Thus, the Center for Tropical Forest Science (CTFS) was born--an informal collaboration of national researchers and their institutions. This became part of the Smithsonian Tropical Research Institute but was managed from the Arnold Arboretum until my retirement in 2000. CTFS has since expanded to become a component of the Smithsonian's new Forest Global Earth Observatory (ForestGEO). This expansion aims to build international capacity in forest science, monitoring the effects of climate change on natural terrestrial ecosystems. The program is now directed by Stuart Davies, who, as one of my Harvard graduate students, completed elegant field observations and experiments on habitat differentiation within a species-diverse genus of pioneer trees, Macaranga, a member of the spurge family (Euphorbiaceae). Crucially, Stuart had already gained the friendship of our regional partners. Although still best represented in East Asia, there are now sixtyseven forest research sites worldwide, including one at Harvard Forest, and more than six million individual trees monitored. More than four hundred published peer-reviewed papers underwrite the massive acceleration in our knowledge of forests. The CTFS focus on understanding tropical rainforest species diversity continues, but the work has revealed unexpected patterns: Plot species diversity, instead of increasing with habitat favorability, unexpectedly peaks at quite low levels of soil nutrients. This, interestingly, supports a theory advanced by ecologist David Tilman, now at the University of Minnesota, in his Princeton doctoral dissertation. He predicted that the low species diversity of plant communities in habitats that are severely lim- ited by low fertility, drought susceptibility, or shadiness is enhanced once these limiting factors start to relax. As soon as these factors relax past a certain threshold, however, one or a few of the species that grow fastest overtop the rest, suppressing subsequent diversity by competitive shading or by hindering establishment. The pattern shown by our plots confirmed the prediction remarkably, independent of the distance between plots. But that is not the whole story. We are still left with insufficient explanation as to how so many tree species can co-occur in a single community. Some years later, our postdoctoral researcher Koichi Kamiya used molecular genetic analysis of seedlings beneath a grove of four distinct but related Shorea species and found that although many were hybrids, very few of the reproductively mature trees were of hybrid origin. This provided clinching evidence that selective mortality results in survival only of those individuals that retain the parental genome and reoccupy the parental, as yet undefined, niche. Competitive selection also leads to differentiation of flowering times, stature, and response to light among sister species, but these explanations are surely insufficient explanations for the co-occurrence of this incredible biodiversity. Joseph Connell, at the University of California, and Dan Janzen, at the University of Pennsylvania, independently proposed that high diversity could be maintained if each species were to attract a single seed predator, such that seed mortality (causing fewer juveniles near parent trunks) would lead to space available for the establishment of others. But no vertebrate seed predators are so specialized. Czech entomologist Vojtech Novotny has convincingly shown, through studies in New Guinea, that herbivorous insects attack at a generic rather than species level. Instead, researchers, notably Yale professor Liza Comita and her students at the Barro Colorado Island plot, have discovered that the prevalent mortality of established seedlings in hyperdiverse rainforest tree communities is mediated by host-specific pathogenic microorganisms, especially fungi and viruses. If mature populations of particular tree species are less dense, the seedlings are less chemically defended. These less-common species include 26 Arnoldia 77\/2 ? November 2019 trees whose seeds and pollen are the most widely dispersed, including species like the wild progenitors of cultivated mangoes (Mangifera), rambutans (Nephelium), and durians (Durio), which produce few fleshy fruits that are sought by mammals and large birds. Now, with hundreds of findings resulting from the CTFS coordination and research continuing to expand, we can conclude that niche specificity does indeed govern floristic structure within and between tree species in hyperdiverse plant communities such as rainforests, except at very local levels where the pull of limited seed dispersal is influential. As such, the pub dispute with Steven Hubbell, back in 1982, can be resolved: ecological niches occupied by particular species become increasingly specialized over time thanks to competitive interactions, so MacArthur and Wilson's theory of island biogeography can generally be applied to rainforest tree biodiversity at the local scale (a habitat island), but the theory rarely applies more broadly, because climate or geological changes ensue before an equilibrium can be reached in the number of large, long-lived plant species that might eventually occupy a nationsized island. When I first began my research career in Borneo, in 1957, the limitless lofty forests, the unforgettable aromas, and the bird-and-cicada orchestra echoing through the cathedral-like subcanopy were nothing short of glorious. Things are now so very different. The two British colonies, Sabah and Sarawak, united as states within independent Malaysia at an ominous time. Peninsular Malaysia had gained independence in 1957, at the moment when the African oil palm (Elaeis guineensis) was beginning to be a serious commercial competitor to the Brazilian rubber tree (Hevea brasiliensis), causing increased demand for new agricultural land. Agronomists in Peninsular Malaysia recommended that all soils that supported mixed dipterocarp forest were suitable for oil palm cultivation. Legal constraints on timber exports were relaxed. The international trade peaked, with oversupply depressing prices. Sustainable forest management languished. Now, Pasoh contains the only inland mixed dipterocarp forest remaining unlogged in Peninsular Malaysia outside the parks. In Borneo, the same fate awaited mixed dipterocarp forests ten years later. With dominance of demand from Japan, and later China, a local wood-based industry, which had benefitted from government investment in research such as mine, went into decline. Brunei, prospering from its oil, has alone retained aboriginal forest over two thirds of its modest land area, timber harvesting being allowed only for the home market. My initial campsite in Brunei, back in 1957, at Kuala Belalong, now hosts a university forest research and training camp, while our plots in the Andulau hills, closer to the coast, are now encompassed within a research preserve and an adjacent forest service research station. In Sabah, thanks in part to political gains among the inland communities, a successful expansion of an ecotourism industry, and an outstanding and farsighted director of forests, Datuk Sam Mannan, large tracts have been conserved and riparian fringes protected. But Sabah was far from immune to events that happened in Peninsular Malaysia and those that followed in Sarawak, where politicians saw timber licenses as a ready bribe to induce candidates to change sides. In Sarawak, politicians and their families became the new rentier elite, with the power to delegate timber licenses, awarded over periods that often corresponded with elections rather than felling cycles of fifty years or longer. Licenses were, in turn, delegated to companies of industrious and enterprising overseas Chinese, who have used their profits to expand their operations as far as New Guinea and South America. Young Dayaks hazarded their lives as saw operators at one hundred dollars a day, too often with tragic consequences. Now, other than in the parks, little of the original rainforest remains in Sarawak. On climbing the basalt peak of Bukit Mersing, in central Sarawak, I recall looking down in wonder on the lavender cascades of the rare strangler Wightia borneensis in flower. But, years later, a silviculturist apologetically confessed to me that this magnificent park, which I had proposed given its rich diversity of rare Eusideroxylon zwageri is a rare tree in the laurel family (Lauraceae), which is listed as vulnerable on the International Union for Conservation of Nature's Red List of Threatened Species. 28 Arnoldia 77\/2 ? November 2019 Oil palm (Elaeis guineensis) cultivation has created a massive conservation threat for mixed dipterocarp forests in Southeast Asia. and endemic species, had \"inadvertently\" been licensed for logging. Our permanent plots there were trashed, and those in coastal Nyabau forest were cleared for an oilfield service depot. Of all our thirteen plot sites in Sarawak, only three remain unfelled. Although smallholder plantations have increased, the relegation of formerly reserved forests to commercial interests has sanctioned a massive transfer of wealth from the rural poor to the new urban rich. Sarawak's national parks and strict preserves, unique among rainforest conservation areas, were selected and demarcated on botanical and ecological criteria rather than mammalian fauna. In the absence of any policy for retaining the animal migration paths, riparian forests between all but two of Sarawak's national parks have been destroyed, and most of the parks are consequently too small to sustain all but populations of the smallest vertebrates. In the summer of 2019, while writing this essay, I received the following from an old Sarawakian friend, Paul Chai, who succeeded me as forest botanist in 1966 and, like me, is long retired: \"No good news on forestry. Sarawak is experiencing annual haze due to burning in Kalimantan and here, and Miri is worse.... I am worried that other national parks may soon be at risk. Enforcement is poor and relies on drones and helicopter.\" This tragedy has resulted from the meteoric rise of China in the international timber trade--a country which has evolved exemplary conservation policies for its own natural resources but which imposes no rules on its overseas commercial interests, including imports of timber, and animals and plants of value for traditional medicine. Yet to whom, ultimately, should the accusing finger be pointed? China now exports more than half the world's furniture, and most of that production is purchased in the West. So, I wonder, have I been wasting my time? Laboring in this depressing environment, though, are two outstanding young Iban Rainforest Biodiversity 29 A mixed dipterocarp forest has been cleared and terraced for oil palm cultivation. women--Julia anak Sang and Wilhelmina anak Cluny, respectively a field botanist and a wildlife naturalist--doing their best to turn the tide with courage and determination. Julia works in the Sarawak Timber Corporation and has a team of forest botany technicians who search the degraded forests for surviving tree flora. She is publishing Red List data for the International Union for Conservation of Nature and is mapping and documenting current species conservation status within and outside the protected areas. Wilhelmina is a conservation officer in the Sarawak Forest Department who has focused on vertebrate conservation. She has worked on enhancing local community involvement in protected area conservation, especially Kayan Mentarang National Park, a large park that spans the border between Sarawak and the Indonesian territory of Kalimantan. Conservation and forestry are not fundamentally incompatible, but given the current status of forest degradation in this region, how long would it take to restore a sustainable felling cycle? Probably fifty to one hundred years. How long to restore the original forest carbon mass? At least one hundred. And species diversity? Even if there is no local extinction, probably at least a millennium. I'm hopeful, at least, that our research and the continued work of CTFS and ForestGEO can provide the information necessary for restoring, in the future, the unimaginable diversity of these rainforests. Reference The map for this article was created using Esri, USGS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, lntermap and the GIS user community. Peter Ashton is Harvard University Bullard Professor Emeritus and was director of the Arnold Arboretum from 1978 to 1987. Among many career honors, his research on tropical forests was recognized with the prestigious Japan Prize in 2007. He and his wife, Mary, live in Chiswick, London. This Arnoldia article is the third and final in a series about Ashton's research career. "},{"has_event_date":0,"type":"arnoldia","title":"E. S. Rogers and the Origins of American Grape Breeding","article_sequence":4,"start_page":30,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25669","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070b328.jpg","volume":77,"issue_number":2,"year":2019,"series":null,"season":null,"authors":"Casscles, J. Stephen","article_content":"CASSCLES, J.S. 2019. E.S. ROGERS AND THE ORIGINS OF AMERICAN GRAPE BREEDING. ARNOLDIA, 77(2): 30?39 E.S. Rogers and the Origins of American Grape Breeding J. Stephen Casscles I n the mid-nineteenth century, eastern Massachusetts was a hub for American horticultural talent, including writers and nursery owners. In the case of fruit breeders like Edward Staniford Rogers, even work conducted at a relatively small scale had the potential to spread nationally, shaping breeding efforts into the present. Rogers focused on grapes, and he did his work in his half-acre backyard at 376 Essex Street, in Salem, Massachusetts. Rogers came from a prosperous mercantile and shipping family, and in 1826, the year Rogers was born, Essex Street was one of Salem's wealthiest residential neighborhoods. The street ran into the city's center, near the wharves of Salem Harbor, then one of the busiest ports on the Atlantic seaboard. Rogers began his breeding work in 1851, when he was still a young man. While shy with most people--some would even say reclusive--he could talk endlessly about his new hybrid grape creations and did so regularly with local and nationally renowned horticulturists. In the process, he became a leading American grape breeder, focusing on hybridizing the common European grapevine (Vitis vinifera) with the hardier and more disease-resistant American fox grape (V. labrusca). Ulysses Prentiss Hedrick, the chief horticulturist of the New York State Agricultural Experiment Station, extensively wrote about Rogers's work and noted, in 1908, that when Rogers introduced his grapes to the public in the late 1860s, \"enthusiasm and speculation ran riot.\" Another breeder, James H. Ricketts of Newburgh, New York, had released successful varieties around the same time, and it was, according to Hedrick, a \"golden era\" for American grapes. \"Possibly at no other period has interest in grape-growing been so keen as during the decade succeeding the introduction of these hybrids,\" he wrote. A Horticultural Renaissance As a grape grower and winemaker, I came to appreciate the landmark hybridization work of Rogers while researching and acquiring cool-climate grape hybrids that were developed in France between 1860 and 1940--primarily crosses between Vitis vinifera and American species like V. labrusca, V. aestivalis, V. riparia, and V. rupestris. I became attuned to these French-American grape hybrids starting in 1974, first by working at Benmarl Vineyards, in Marlboro, New York, and subsequently at the Hudson-Chatham Winery in Ghent, New York, where I encountered red grapes like `Baco Noir' and `Chelois' and white grapes like `Seyval Blanc' and `Vidal', along with more than a dozen others. My interest then led me to Rogers and other East Coast hybridizers, including a significant number based in the Hudson Valley. I evaluated the Rogers varieties to see if they could be grown in a more ecologically sustainable manner than grape varieties that are conventionally grown today, and I wanted to learn more about the flavor profiles of these forgotten old varieties. For over fifteen years, I have grown twelve of these Rogers hybrids very successfully on my farm, Cedar Cliff, in Athens, New York, for wine production. It is my hope to reintroduce some of the Rogers grapes to commercial growers and wineries so that they can be made more readily available to the public. In the process of researching heirloom grape varieties, I discovered that, between 1840 and 1890, eastern Massachusetts and the MidHudson Valley were two of three centers of grape breeding. Breeders were also busy near St. Louis, Missouri. In Massachusetts, the breeders were generally wealthy New England Brahmins, like Rogers, whose families made fortunes either as merchants or in mercantile shipping. Edward Staniford Rogers began hybridizing American and European grapes in 1851, launching an era of \"enthusiasm and speculation\" for American grape growers. All images, unless indicated, are from The Grapes of New York by Ulysses Prentiss Hedrick. ALL IMAGES FROM BIODIVERSITY HERITAGE LIBRARY 32 Arnoldia 77\/2 ? November 2019 The American fox grape (Vitis labrusca, left) is well-known for being a parent of the spontaneous hybrid `Concord' (right), which is often used for jellies and juices. These genteel farmers engaged in horticulture for intellectual stimulation and social comradery. Nursery owner Charles Mason Hovey facilitated communication between nearby breeders and regularly corresponded with the national fruit-breeding community either in person or through the many plant catalogues, pamphlets, and horticultural books that he wrote. In addition, Hovey helped to direct, along with Marshall Pinckney Wilder, the nationally recognized Massachusetts Horticultural Society and the American Pomological Society. Local agricultural and horticultural societies were also actively evaluating new horticultural varieties in most eastern Massachusetts counties. Several forces compelled these Massachusetts horticulturists to develop new hybrid fruit. Many desired to create plant material for their suburban and rural homes, emulating the British landed gentry and securing greater social prestige within their community. Oth- ers desired to develop fruits that were more productive and disease resistant for profit. An underlying theme was the uniquely New England quasi-religious-social-ethical belief among business, social, and religious leaders that one's religious service could be manifest by service to community. Work had a moral component, and the highest calling was to be productive; unlike the trading of goods, engaging in agriculture and manufacturing was a divine calling. Further, by 1800, the region's already thin agricultural soils were becoming very depleted due to more than a century of extensive but unwise cultivation techniques and practices. Hence, a movement arose to study agriculture, hybridization, and plant sciences, so that local farmers could revitalize their increasingly poor and overcropped soils. The business community supported these agricultural research initiatives so that farmers would remain in Massachusetts and continue to be their loyal American Grape Breeding 33 Rogers made careful crosses between the fox grape and the European wine grape, Vitis vinifera (left). He used a common variety known as `Black Hamburg' (right). customers, instead of being forced to move farther west in search of more fertile soils. The business community's support was evidenced by the founding of the Massachusetts Society for Promoting Agriculture in 1792. Its membership was clearly mercantile in composition, including most of eastern Massachusetts's prominent families, along with attorneys and a few physicians and clergy, most of whom were Harvard College alumni. Hybrid Crosses Even within this vibrant horticultural milieu, Rogers was unique. According to Thomas Volney Munson, a central figure among the next generation of American grape breeders, Rogers was responsible for taking \"the first intelligent step\" towards developing \"thorobred\" American grape varieties. Unlike the classic `Concord', which was selected by Ephraim W. Bull from a spontaneous cross between Vitis labrusca and V. vinifera (the results of natural insect pollination) in Concord, Massachusetts, in the 1840s, Rogers was intent on carefully making and documenting his crosses. In his own words, Rogers said, \"When I commenced experimenting I had no knowledge of any one who had raised grapes by this process, though I had heard of flowers, pears, &c., and I had attempted crosses of pears. Reading articles in the London Horticulturalist, it occurred to me that I could get a new grape by this process; combining the qualities needed for open culture, it would be more valuable than any other fruit.\" Rogers was drawn to the quiet and contemplative life of horticulture, and once his father died in 1858, he very quickly exited the family shipping business and concentrated on his horticultural pursuits and real estate investments in Rockport, Massachusetts. He wanted to create new grape varieties that incorporated 34 Arnoldia 77\/2 ? November 2019 the more sophisticated and subtle flavors of European Vitis vinifera varieties (like the table grapes we buy today in the supermarket) with the hardiness and reliable productivity of native American grape varieties, ripening early, before the first fall frost. Successful varieties also needed to possess ample fungal disease resistance and simultaneously be productive enough as commercial table grapes, with big berries, big clusters, sufficient sweetness, and skin that adhered to the flesh of the berry. In the summer of 1851, Rogers made crosses using a seed parent, Vitis labrusca `Carter' (a wild-type variety also known as `Mammoth Globe'), and the pollen of V. vinifera `Black Hamburg' and `White Chasselas'. `Carter' was used as the seed parent because this self-sterile variety was large fruited, hardy in the field, and one of the earliest ripening local selections that he could find. The pollen of `Black Hamburg' and `White Chasselas' was chosen because they were two of the hardiest European varieties and were the most commonly available in Massachusetts. The pollen was obtained from vines growing in a nearby unheated glass greenhouse. The exact provenance is unrecorded, but the pollen could have come from someone like John Fisk Allen, who lived about two blocks from Rogers, or George Haskell, in nearby Ipswich. Both men were highly interested in grape cultivation. The `Carter' blossoms were emasculated and fertilized with Vitis vinifera pollen and small cotton bags were placed over the `Carter' female flowers. Rogers also placed clusters of V. vinifera blossoms in the bags. From this cross-pollination, he secured about 150 seeds. These seeds were then planted in his backyard garden that fall. The following spring, many of these seeds germinated, but cut worms and other accidents reduced the number of vines to forty-five. These forty-five vines grew upward on poles for three years. Due to overcrowding, Rogers transplanted twenty-five of the plants to other parts of the garden to give them enough room to grow. The untransplanted vines started to bear fruit in 1856, and the transplanted varieties fruited a few years later. In observing the garden, Marshall Pinckney Wilder, of the American Pomological Society, said, \"How much can be done with little is illustrated by the fact that all [of his grapes] ... were produced by a lame man in a half-acre city lot 150 years in cultivation.\" Further, he noted that the lot was \"a cold matted soil filled with old apple and pear trees, currant bushes, flax and everything mingled in together.\" Rogers believed his grape creations to be a success, noting the intermixture of traits between the species. \"The vines are even more vigorous than the parents,\" he wrote, \"and more exempt from diseases, and more hardy than most outdoor varieties.\" The seedlings were numbered one to forty-five. In 1858 and 1859, Rogers sent cuttings of these numbered varieties to growers and horticulturists for further testing. He disseminated these varieties due to the small size of his backyard garden and because the common practice then, as it is now, was to share plant material with colleagues to get comments on the growing attributes, strengths, and weaknesses of such plants in a wide range of climates and soil types. Through his painstaking work, Rogers created over twenty major grape hybrids. The resultant grapes were first officially introduced to the public in 1867. In 1869, Rogers named thirteen of his varieties after local Massachusetts places and people (`Agawam', `Massasoit', `Salem', `Essex', and `Merrimac'), as well as for horticulturists (`Barry', `Lindley', `Gaertner', and `Wilder') and the German writer Johann Wolfgang von Goethe (`Goethe'). These were promoted through the Catalogue of Fruits by the American Pomological Society, an organization that was based in Boston. From there, the Rogers hybrids steadily gained interest and notoriety across the United States and Canada. The Rogers Grapes All Rogers hybrids possess large or very large berries, medium-sized clusters, and grape skin that is either attached or semi-attached to the berry flesh, unlike the \"slip-skin\" characteristic of the `Concord'. They grow vigorously, have better fungal disease resistance than their European pollen parents, and are hardy and American Grape Breeding 35 By 1869, Bushberg Vineyards and Orchards, in Missouri, promoted many of the Rogers grape varieties in their Illustrated Descriptive Catalogue of Grape Vines, Small Fruits, and Seed Potatoes. Of `Goethe', the catalogue advertised, \"At the fall meeting of the Mississippi Valley Grape Growers' Association, September 9, 1868, we exhibited for the first time a few branches of the vine, each with several perfect clusters, which were much admired, and would have probably astonished even its originator, could he have seen them.\" 36 Arnoldia 77\/2 ? November 2019 productive. I like the growing characteristics of the twelve Rogers hybrids that I cultivate in the Mid-Hudson Valley, and the resultant fruit is flavorful and makes wonderful wines that have an attractive combination of soft flavors of Muscat grapes and Vitis labrusca. These characteristics made the Rogers hybrids very popular when first introduced to America and Canada in 1867. They were initially quite sought after by growers, talked about at horticultural and agricultural society meetings, and widely evaluated. In 1895, the nationally recognized Bushberg Vineyards catalogue, which set the standard for fruit catalogues and pomological literature in North America, extensively covered the Rogers hybrids with accompanying illustrations of many of them. The Bushberg catalogue stated that these Rogers varieties were \"very productive,\" \"beautiful,\" and \"valuable\" selections that were \"handsome in appearance\" and of \"fine quality\" for the table and for wine. Other definitive North American nursery catalogues of the latter nineteenth century, including Hovey's The Magazine of Horticulture, prominently featured and illustrated the Rogers hybrids, as did agricultural magazines like The Gardner's Monthly and Horticulturist, The Rural New Yorker, and The Country Gentleman. Among Rogers's selections, `Agawam' is one of his best. In 1908, Hedrick reported that `Agawam' was the most widely grown of the Rogers hybrids, noting that it was sold by practically all nurseries in the United States east of the Rocky Mountains. It is the only completely self-fertile of the Rogers varieties. The color is a dark purplish-red with a lilac bloom. The wines are aromatic with rich fruit flavors of Muscat grapes and hints of fresh grapes, guava, and tropical fruits from Vitis labrusca, along with an herbal finish. The body is substantial and viscous for a white wine, and it can either stand alone or be used in blends with other white wines. Tasting something like this is to taste the nineteenth-century innovation of Rogers and his contemporary fruit breeders in Massachusetts, the Mid-Hudson Valley, and the St. Louis area. The Rogers Hybrids Live On A combination of factors led to dwindling name recognition for the Rogers grapes. Hedrick stated that the period between 1853 (the date `Concord' was first introduced) and 1880 could be \"singled out as the period in which viticulture made its great growth in eastern America.\" After 1880, however, California started to compete in earnest with eastern vineyards, and grape prices fell significantly in eastern metropolitan markets, given the vast influx of inexpensive California grapes. This competition, combined with higher incidence of fungal diseases and insect damage in eastern vineyards, which were planted too closely to one another, severely reduced overall production in the east. With a corresponding reduction of grape acreage, varieties like `Concord', `Niagara', and `Delaware' expanded their dominance, while the Rogers hybrids, which, save for `Agawam', were mostly self-infertile, declined relatively and absolutely in acreage. In addition, the enactment of Prohibition in 1920 further reduced their demand for wine production, which was their primary use. Yet the Rogers hybrids live on in the twentyfirst century. In the Rogers era, privately organized horticultural and agricultural societies, such as the Massachusetts Horticultural Society, sponsored the bulk of the public discussion about plant evaluation. However, with the Congressional enactment of the Morrill Act of 1862 (establishing agricultural land-grant colleges) and the Hatch Act of 1887 (establishing agricultural experiment stations), this horticultural domain shifted increasingly to government-financed programs. With this shift, many of the Rogers grapes were incorporated into the most advanced American cool-climate, wine-grape breeding programs of the twentieth century. For example, at Cornell University, the Rogers hybrid `Herbert' was used to breed the hybrids `Sheridan', in 1921, and `Buffalo', in 1938. These, in turn, lead to the development of twenty-first-century introductions like `Geneva Red', `Corot Noir', and `Noiret'. Elmer Swenson of Wisconsin, whose private breeding program was subsequently absorbed into research at the University of Minnesota, used American Grape Breeding 37 The author considers `Agawam' to be the finest Rogers variety for modern winemaking. 38 Arnoldia 77\/2 ? November 2019 While the fame of the Rogers grapes waned in the early twentieth century, `Goethe' (left) has found unexpected popularity in Brazil. Also shown: `Lindley'. the Rogers variety `Wilder' as a great grandparent to create `Marquette'. The varieties `Marquette' and `Noiret' are now finding their place in today's North American cool-climate wine industry. In addition, Rogers varieties were used in the breeding programs at agricultural experiment stations in Missouri and South Dakota. Thomas Volney Munson, the pioneer American grape breeder who privately bred many scores of high-quality hardy and disease-resistant grape varieties for the central and southern United Sates, relied heavily on Rogers hybrids for his extensive breeding program. The Rogers varieties do not simply persist as the basis for subsequent breeding efforts. The variety `Goethe' is the foundation of one niche segment of the Brazilian wine industry in the Urussanga region of the state of Santa Catarina. In Brazil, `Goethe' is made mostly into sparkling wines with vineyards that cover over one hundred acres. While `Goethe' is traditionally a pink- red variety, a natural mutation, first observed in a Brazilian vineyard the 1950s, has produced a white clone, now known as `Goethe Primo'. This new variety makes still and sparkling wines that are very Vitis vinifera-like in their flavor profile and acid balance but with pleasant, soft aromatics from V. labrusca. In this region, over twenty thousand gallons of `Goethe' wine are produced, with the remainder sold as table grapes. Today, commercial and hobbyist growers, foodies, farm-to-table advocates, private grape breeders, and university breeding and agricultural research programs are all looking for the \"next best\" fruit variety that is flavorful and productive and which can be grown in a more environmentally sustainable manner. The Rogers hybrids, along with other heirlooms bred in New England and in the Hudson Valley, fit that bill. Rogers's work demonstrates that sometimes the search for the \"next best\" may involve looking back. American Grape Breeding 39 References Bush & Son & Meissner, Bushberg Vineyards. 1895. Illustrated descriptive catalogue of American grape vines (4th ed.). St. Louis: R.P. Studley and Co. Casscles, J.S. 2015. Grapes of the Hudson Valley and other cool climate regions of the United States and Canada. Coxsackie, NY: Flint Mine Press. Cedar Cliff Nursery. 2013?2019. Cedar Cliff Nursery: Grape vines for grape growers and field & cellar notes. Catskill, NY: Casscles, J.S. Downing, A.J., and Downing, C. 1869. The fruits and fruit trees of America (2nd ed.). New York: John Wiley & Sons. Hedrick, U.P. 1908. The grapes of New York (Annual report of the State of New York, Department of Agriculture; no. 15, v. 3, pt. 2). Albany, NY: J.B. Lyon Company. Hedrick, U.P. 1933. A history of agriculture in the State of New York (The New York State Agricultural Society). Albany, NY: J.B. Lyon Company. Hedrick, U.P. 1950. A history of horticulture in America to 1860. New York: Oxford University Press. Hedrick, U.P. 1919. Manual of American grape-growing. New York: The Macmillan Company. Hovey, C.M. The magazine of horticulture, botany, and all useful discoveries and improvements in rural affairs. Boston: Hovey and Co. Hutchinson, B.J. 1980. A taste for horticulture. Arnoldia, 40(1): 31?48. McLeRoy, S., and Renfro, R.E. 2008. Grape man of Texas: Thomas Volney Munson and the origins of American viticulture. San Francisco: The Wine Appreciation Guild. Morton, L.T. 1985. Winegrowing in eastern America: An illustrated guide to viniculture east of the Rockies. Ithaca, NY: Cornell University Press. Munson, T.V. 1909. Foundations of American grape culture. New York: Orange Judd Company. Parker, S.J. 1865. Improvement of native grapes by seedlings and hybridization. In Report of the Commissioner of Agriculture for the Year 1864 (pp. 122?136). Washington, DC: Government Printing Office. Rogers Family Papers, Peabody Essex Museum (MSS 87). Phillips Library at the Peabody Essex Museum, Salem, Massachusetts. Schofield, E.A. 1988. \"He sowed; others reaped\": Ephraim Wales Bull and the origins of the `Concord' grape. Arnoldia, 48(4): 4?15. Thorton, T.P. 1989. Cultivating gentlemen: The meaning of country life among the Boston elite, 1785? 1860. New Haven, CT: Yale University Press. Wagner, P.M. 1969. American wines and wine-making. New York: Alfred A. Knopf. J. Stephen Casscles comes from a fruit-growing family rooted in the Hudson Valley since the 1870s. In 1990, he established a four-acre vineyard, Cedar Cliff, in Athens, New York, where he has concentrated on identifying, growing, evaluating, and propagating heirloom grape varieties that were first developed in New York in the mid-nineteenth century. He has been the winemaker at Hudson-Chatham Winery, in Ghent, New York, since 2008. In 2015, he published a book on historic grape cultivation titled Grapes of the Hudson Valley and Other Cool Climate Regions of the United States and Canada with Flint Mine Press. 36673667 U.S. POSTAL SERVICESTATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION(Required by 39 U.S.C. 3685) 1. Publication Title: Arnoldia. 2. Publication No: 0004?2633. 3. Filing Date: October 1, 2019. 4. Issue Frequency: Quarterly. 5. No. of Issues Published Annually: 4. 6. Annual Subscription Price: $20.00 domestic; $25.00 foreign. 7. Complete Mailing Address of Known Office of Publication: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 8. Complete Mailing Address of Headquarters of General Business Office of Publisher: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 9. Full Names and Complete Mailing Address of Publisher, Editor, and Managing Editor: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500, publisher; Jonathan Damery, Arnold Arboretum, 125 Arborway, Boston, MA 02130?3500, associate editor. 10. Owner: The Arnold Arboretum of Harvard University, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 11. Known Bondholders, Mortgagees, and Other Security Holders Owning or Holding 1 Percent or More of Total Amount of Bonds, Mortgages, or Other Securities: none. 12. The purpose, function, and nonprofit status of this organization and the exempt status for federal income tax purposes have not changed during the preceding 12 months. 13. Publication Name: Arnoldia. 14. Issue Date for Circulation Data Below: September 5, 2019. 15. Extent and Nature of Circulation. a. Total No. Copies. Average No. Copies Each Issue During Preceding 12 Months: 1,600. Actual No. Copies of Single Issue Published Nearest to Filing Date: 1,600. b. 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I certify that all information furnished on this form is true and complete. Jonathan Damery, Associate Editor. "},{"has_event_date":0,"type":"arnoldia","title":"A Teacher's Favorite: Gleditsia aquatica","article_sequence":5,"start_page":40,"end_page":41,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25668","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070af6f.jpg","volume":77,"issue_number":2,"year":2019,"series":null,"season":null,"authors":"Caballero McGuire, Ana Maria","article_content":"MCGUIRE, A.M.C. 2019. A TEACHER'S FAVORITE: GLEDITSIA AQUATICA. ARNOLDIA, 77(2): 40 A Teacher's Favorite: Gleditsia aquatica Ana Maria Caballero McGuire E veryone knows that teachers should not have favorites, but I do. My favorite has muddy feet, a thorny disposition, and reddish-brown, almond-shaped eyes. Oh, and he's also a southerner. Gleditisa aquatica (accession 201-93*B), also known as the water or swamp locust, is a North American native, closely related to the more familiar honey locust (G. tricanthos). You will find the species growing along riverbanks and marshes in its natural range, stretching from South Carolina to central Florida, across Louisiana to eastern Texas, and up the Mississippi River valley to southern Illinois and Indiana. My Arboretum favorite was wild collected in southeastern Missouri. When I introduce children to this tree at the Arboretum, I often start with, \"Who here is brave, really brave? I want to show you a dangerous plant.\" That usually elicits excitement and a loud chorus of \"Me!\" I bring them to Rehder Pond, where they stand looking very closely at the tree behind me. It can take a minute before they understand what they are looking at: a profusion of three- to five-inch-long reddish-brown thorns growing both on the lower parts of the trunk and out along the branches. I often clip a sample and model how to use a one-finger touch along the edge of the thorn to compare its smoothness with the sharp prickly point. It doesn't take long before many children begin to touch the thorns and even ask if they can hold it. They remind me of times when my brothers and I would beg our parents to give us their plastic sword cocktail picks, and we would sword fight in the restaurant while waiting for our meals! Once the children are comfortable with the thorns, we begin a conversation around function. Why would this tree have such thorns? Students quickly identify defense as the main function but then are stumped when asked what the tree is defending itself from. The most common answer is people and predators like foxes, lions, and sharks. It takes some pretend modeling of large herbivores eating before children understand how this tree, having large, thick, and sharp thorns growing at the base of each bipinnately compound leaf might deter large mammals--whether living (deer) or extinct (mastodon)--from eating the leaves. One season I noticed that an American robin (Turdus migratorius) had built a nest on a low branch. In the nest were three young chicks, and the momma was busy flying back and forth, attending to their needs. I used this opportunity to continue the thorn discussion by posing a new question: \"Is that robin smart for building a nest in this thorny tree?\" The group was evenly split between yes and no. Each child had to state their opinion and provide a reason for their answer. In this way, I encourage children to take what they know and what they observe firsthand to form a more complete understanding of how nature works. They also learn to debate by listening to differing views. Aside from thorns, G. aquatica also produces curious eye-shaped seedpods, about 1.5-inches long and flat. Before the seedpod dries out and turns a rich caramel brown color, children can raise the fruit to the sky and see through the papery thin walls to the singular round seed in the middle. Two of these seedpods placed over my eyes elicits cries of \"Owl eyes!\" This fruit is unique among all Gleditsia species because it does not contain a sticky, honey-like pulp surrounding the seeds, and it usually has one seed, rather than ten or more like the honey locust. This difference has led some botanists to suggest that G. aquatica evolved to disperse its seeds via water, instead of animal digestion. Finally, how can I resist a quick math lesson when observing the leaves? The compound leaves measure up to thirty inches long, and the small leaflets occur in six to fourteen pairs on a leaf. They are perfect for a lesson about odd and even and help facilitate counting by twos. Later, children line up in pairs, just like their leaf, and count by twos as they slowly walk back to their bus, heads full of wonder and a pocket or two hiding large owl eyes. Ana Maria Caballero McGuire is the nature education specialist at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23467","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14e816e.jpg","title":"2019-77-2","volume":77,"issue_number":2,"year":2019,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Valuing Biodiversity","article_sequence":1,"start_page":2,"end_page":15,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25667","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070ab6b.jpg","volume":77,"issue_number":1,"year":2019,"series":null,"season":null,"authors":"Grossman, Jake J.","article_content":"GROSSMAN, J.J. 2019. VALUING BIODIVERSITY. ARNOLDIA, 77(1): 2?15 Valuing Biodiversity Jake J. Grossman L ying on my side, I inspect each leaf of a four-year-old red maple (Acer rubrum) sapling. It is midsummer in central Minnesota, and, despite the hot weather, the leaves of this tree and most of its neighbors are covered in brown spots. These spots are a symptom of maple anthracnose, which is caused by infection with any of several ascomycete fungi. Especially in small trees, like the one I am inspecting, anthracnose can slow growth by impairing photosynthesis in damaged leaves and make them more vulnerable to other infections. I finish grading the infected leaves on my tree of interest, record the measurements on my clipboard, and roll over to check its nearest neighbor, also a red maple. Planted only 50 cm (about a foot and a half) apart from each other, these trees are part of the Forests and Biodiversity (FAB) experiment at the Cedar Creek Ecosystem Science Reserve, a long-term research site funded by the National Science Foundation. These trees are in a red maple monoculture plot, meaning each tree has only other red maples as neighbors. Many of these trees are covered in leaf anthracnose, probably because the fungal pathogens that cause it overwinter in the layer of dead leaves, or litter, of infected trees and emerge in the spring to reinfect new growth. Having so many red maples around seems to make any given tree more likely to develop anthracnose. But when I stand up and walk only a few feet away from this maple monoculture, which only reaches my mid-thigh after several years of growth, I encounter a markedly different part of the FAB experiment. On entering a \"biculture,\" or two-species plot, of red maple and jack pine (Pinus banksiana), I am surrounded by pines that exceed my own height and maples that reach my chest. These trees were planted at the same time as the maples in the adjacent monocultural plot, but living with diverse neighbors has clearly made a difference to them. Whereas maples in monoculture tend to be short and stocky, with little space between leaf buds, maples that are forced to compete for light with faster-growing pine tend to come up long and spindly, with tons of space between each set of leaf buds. And, exposed to a lower level of accumulated leaf litter from other red maples, these trees have a much lower incidence of leaf anthracnose. These two plots illustrate two extremes of the FAB experiment, which I comanaged while a graduate student at the University of Minnesota. Previously, as a young ecologist, I had devoured the reports of experiments designed to assess the importance of biodiversity coming out of Cedar Creek and applied to a doctoral program at Minnesota in the hopes that I would get to work there. I quickly heard back from Jeannine Cavender-Bares, a plant ecophysiologist and evolutionary ecologist who would become my doctoral advisor. At the time, she and a group of colleagues at Minnesota were in the final stages of planning a tree biodiversity experiment designed to expand on the foundation laid by grassland experiments at Cedar Creek and across Europe. This group was open to bringing in a new graduate student to help with the establishment of the new project and to do some preliminary research. Freshly returned from two years as a Peace Corps volunteer in subtropical Paraguay, I dug out my long underwear and moved to Minneapolis. On the Origin of Biodiversity Research But why go to the trouble of planting thousands of trees in various combinations of species, then take the time to make thousands of minute measurements of their every centimeter of growth and bout with an illness or pest? For me, as for a generation of researchers at Cedar Creek, experiments like FAB have emerged as a powerful approach for asking what role biodiversity, meaning the variety of life in a particular place or across the globe, might play in keeping the natural world working in the way PHOTOS BY AUTHOR UNLESS NOTED Biodiversity Research 3 To understand whether more-biodiverse tree communities are more productive than less-biodiverse communities, Jake Grossman and colleagues at the University of Minnesota designed research plots with different combinations of tree species, including a two-species biculture (top left) and a five-species polyculture (top right). Grossman is pictured within one of the five-year-old polyculture plots. JACOB MILLER, CEDAR CREEK ECOSYSTEM SCIENCE RESERVE (CC BY-SA 4.0) 4 Arnoldia 77\/1 ? August 2019 Over the last thirty years--a time of increasing global concern about the consequences of global diversity loss--ecologists, as well as farmers, foresters, conservationists, economists, and policymakers, have begun to ask how we measure the extrinsic value of biodiversity. we prefer it to do. To understand these experiments and the findings that come from them, though, we ought to take a step back and consider the history of what I would call the science of biodiversity. When we speak of the environmental challenges of the current era, we can hardly avoid worrying about the erosion of biodiversity. While considerable disagreement persists over how biodiversity across the earth's diverse landscapes has changed over the last ten thousand years, a period in which humans have exerted a growing influence over the biosphere, a consensus has emerged that, at a global level, our planet has entered into a period of precipitous biodiversity loss (Butchart et al., 2010). A case in point: a recent report from a group of Danish and Swedish authors predicts that the loss of mammal diversity that has taken place since the end of the last Ice Age will take 2.5 billion years--two-thirds of the time during which there has been life on the planet--to be replenished by natural evolutionary processes (Davis et al., 2018). Yet such sobering statistics beg a second question: does biodiversity loss really matter? Of course, to many of us--including, I imagine, most readers of Arnoldia--the diversity of earth's species represents an irreplaceable gift. We sense a precious value, whether spiritual, emotional, or cultural, inherent to the diversity of life on earth. It is challenging, though, to convince others. And so, those of us who wish to protect biodiversity must ask ourselves whether there is an extrinsic value to diversity and, if so, how we can justify its conservation. The ecosystem services movement has answered this question by, at least to some extent, evaluating biodiversity in terms of dollars and cents. For instance, Canadian scholars Robin Naidoo and Wiktor Adamowicz (2006) estimate that the financial returns from visits by ecotourists to a Ugandan park rich in bird biodiversity far exceed the costs of maintaining the Biodiversity Research 5 park. This approach, however, doesn't fully capture a deeper question: does biodiversity support the vast array of ecosystem processes-- or functions--that keep our biosphere working and, in doing so, sustain human life? In other words, are more-biodiverse ecosystems stronger and more resilient? The history of this question is a long one, with origins prior to the formal elaboration of the concepts of biodiversity (by American conservation biologist Raymond F. Dasmann in 1968), of ecosystems (by English botanist Arthur Tansley in 1935), and of ecology itself (by German biologist Ernst Haeckel in 1866). Instead, the question of how biodiversity affects ecosystem function was posed first, at least within Western scientific discourse, by the founding mind of modern biology, Charles Darwin. While Darwin is known foremost for his role in developing the concept of evolution by natural selection, his works also offer up a clairvoyant catalogue of research questions for contemporary biologists, one that we have yet to plumb fully some 130 years following his death. Through his lifetime, Darwin contributed important insights to the study of insect pollination; plant physiology; soil formation; the genetic origins of animal behavior; and the natural history of barnacles, coral reefs, and carnivorous plants; among other topics. Indeed, if we turn to Darwin's On The Origin of Species, first published in 1859, we find a claim that, though peripheral to the broader case for adaptive evolution, constitutes the origin of an important field of biodiversity research: \"If a plot of ground be sown with one species of grass, and a similar plot be sown with several distinct genera of grasses, a greater number of plants and a greater weight of dry herbage can thus be raised.\" In this brief aside, Darwin argues that it was, at the time, well known that more-biodiverse systems--well, grasslands, at least--ought to be more productive than less-diverse ones. Indeed, indigenous peoples, and especially farmers, have known for millennia that morediverse ecosystems are more productive. For instance, the \"three sisters\" technique of growing diverse gardens of corn, beans, and squash developed in pre-Columbian central Mexico and subsequently radiated throughout the Americas. Contemporary studies have demonstrated that this system of polyculture--or growing multiple crop species together--boosts yield compared to monocultures of constituent species (Zhang et al., 2014). Experimental assessment of traditional Chinese polycultures consisting of varying mixtures of wheat, corn, and soybeans have revealed similar trends (Zhang and Li, 2003). Such traditional techniques have continued to evolve to this day, resulting in, among other practices, the contemporary interest in \"companion planting\" among home gardeners and farmers. For instance, many gardeners in North America are familiar with the practice of planting African marigolds, mints, and other aromatics in their gardens, both for their own aesthetic or culinary uses and, allegedly, to deter pests. Despite this mountain of traditional knowledge and practical evidence, the empirical reality of the link between biodiversity and ecosystem function went without formal evaluation for over a century before slowly climbing back into the crosshairs of experimental biologists. An Ecological Reawakening For much of the twentieth century, ecologists explored tantalizingly around the question of how biodiversity might shape ecosystems, often taking diversity to be a consequence of ecological conditions in a particular place rather than a cause of those same conditions (e.g., Connell and Orias, 1964). Eventually, as ecologists became more attuned to the ecological importance of stability--how much conditions in a forest or grassland might remain constant from season to season or year to year-- they began to interrogate its relationship with biodiversity. At the center of this debate was the question of whether increasing the number of species in a community made that community more stable through beneficial effects such as symbiosis (Elton, 1958) or destabilized it by increasing the likelihood of, for instance, local numbers of a critical species crashing due to catastrophic disease (May, 1973). In their review of the field, American ecologist David Tilman and colleagues (2014) trace a \"reawakening\" in the study of biodiversity 6 Arnoldia 77\/1 ? August 2019 to one of three biodiversity treatments. The lowest diversity rooms contained boxes of soil enriched with two common British plant species (e.g., sow thistle), three plant-eating invertebrates (e.g., aphids), one predator (e.g., an aphid predator), and three decomposer species (e.g., earthworms). A second set of rooms contained extra species of each class, and the most diverse rooms contained sixteen plants, five herbivores, two predators, and eight decomposers. Environmental conditions were held constant, and during a two-hundred-day period, an international team of ecologists monitored a variety of ecosystem functions: how much organisms in each room respired, how quickly organic matter decomposed, to what extent nutrients and water ran off, and how productive plants were in each room. In the end, morediverse communities of plants and animals consumed more carbon dioxide (respired more) and grew more than less-diverse ones (Naeem et al., 1994). Diversity supercharged the functionality of ecosystems with more species. JACOB MILLER, CEDAR CREEK ECOSYSTEM SCIENCE RESERVE (CC BY-SA 4.0) to incipient awareness of catastrophic global biodiversity loss during the 1980s, which culminated in a 1991 conference of ecologists in Bayreuth, Germany. The papers emerging from this meeting--which were ultimately collected in an edited volume, Biodiversity and Ecosystem Function, published in 1994--effectively launched the field of contemporary research on biodiversity-ecosystem functioning, otherwise known as BEF. The observational findings and theories marshaled in the very early nineties, however, lacked the gold standard of ecological evidence: experimentation. This was not long in coming; two progenitor BEF experiments were already in development at the time of the Bayreuth Conference. At Imperial College London's Centre for Population Biology, pilot testing of the futuristically named Ecotron facility began in 1991. The Ecotron, still operational today, consists of sixteen isolated rooms, each with its own light, temperature, and atmospheric control systems. Beginning in 1993, these rooms were assigned Between 1994 and 1995, researchers at the Cedar Creek Ecosystem Science Reserve established the \"Big Biodiversity\" experiment, which was one of the first field-based experiments to provide empirical evidence about the relationship between biodiversity and ecosystem function. Biodiversity Research 7 efficiently utilized available nutrients (Tilman et al., 1996). These findings have been borne out repeatedly through the expanded \"Big Biodiversity\" experiment, planted between 1994 and 1995. These plots are larger, more numerous, and contain as many as thirty-two species (Tilman et al., 1997). The assigned diversity levels of most of its original plots are still maintained through diligent weeding by an army of freshfaced interns hired by Cedar Creek's managers every summer. Now in its twenty-fifth year of growth, the Big Biodiversity experiment still serves as a critical platform for BEF research. A FABulous Journey The Forests and Biodiversity (FAB) project, which I was recruited to work on in 2012, would mimic past grassland experiments insofar as plots were planted with varying species diversity. Yet, in many other ways, the forest project departed from its progenitors. From a logistical standpoint, rather than weighing out JACOB MILLER, CEDAR CREEK ECOSYSTEM SCIENCE RESERVE (CC BY-SA 4.0) In a complement to the highly controlled approach of the Ecotron, Tilman and collaborators at Cedar Creek, in Minnesota, were simultaneously figuring out how to ask BEF questions in the field. Based on an observational study in which more-diverse grassland plots showed greater stability in biomass production than less-diverse plots following an extreme drought (Downing and Tilman, 1994), they established what came to be known as Cedar Creek's \"Little Biodiversity\" experiment. In this seminal experiment, 147 plots, each nine meters square, were denuded of existing vegetation and seeded with one, two, four, six, eight, twelve, or twenty-four species of prairie plants. Echoing findings from the Ecotron, diverse plots (and especially any plot with twelve or twentyfour species) produced far more biomass than less-diverse plots. Furthermore, even after only two summers of growth, more-diverse plant communities in the Little Biodiversity experiment showed lower levels of soil nitrogen, suggesting that their roots more completely and Researchers have expanded on the \"Big Biodiversity\" methods to explore the relationship between biodiversity and climate change at additional prairie plots. 8 Arnoldia 77\/1 ? August 2019 and broadcasting consistent quantities of seed, we planted each tree on a grid (sixty-four trees per plot), where each tree was only half a meter from its nearest neighbors. Some plots were monocultures, consisting entirely of one of twelve species native to Minnesota: red (Pinus resinosa), jack (P. banksiana), or white pine (P. strobus); eastern red cedar (Juniperus virginiana); paper birch (Betula papyrifera); red (Quercus rubra), northern pin (Q. ellipsoidalis), bur (Q. macrocarpa), and white oak (Q. alba); basswood (T. americana); red maple (Acer rubrum); and box elder (A. negundo). Other plots (bicultures) contained thirty-two individuals of each of two species. Yet others were planted with five-species polycultures, and we threw the entire kitchen sink at a set of twelve-species plots. We started with twoyear-old bareroot seedlings, planted in May of 2013, and over the next three years, we replanted dead trees and weeded woody invaders so that each plot truly corresponded to its assigned tree-diversity treatment. By the time I finished my doctorate five years later, I could easily conceal myself within their densely interlocking boughs--at least in plots dominated by fast-growing pines and birches. Beyond logistical considerations, the design of FAB also expanded on past research by making it possible for us to ask which dimensions of biodiversity might be most important to supporting ecological function. For instance, the vaunted boost in productivity associated with higher-diversity plots in the Big Biodiversity grassland study appears not to be entirely due to species richness--the number of species in a plot. Instead, it seems that some of the diversity-related boost really stemmed from functional diversity, the variability in morphological and physiological traits associated with The Forests and Biodiversity (FAB) plots were established in 2013, with sixty-four seedlings planted in different combinations within each plot--totaling 140 plots and almost nine thousand trees. Biodiversity Research 9 species in a community. In particular, it appears that more-diverse plots provided opportunities for nitrogen-fixing legumes and droughttolerant grasses to interact synergistically, boosting the productivity of their community by sharing resources. Legumes fertilized nearby grasses, which, because they differ in their growth form and resource needs, did not outcompete their beneficial neighbors (Fargione et al., 2007). In this sense, it can sometimes be difficult to determine whether it is more important to have a lot of species present or for those species present to have a diversity of functions. While functional diversity can be difficult to measure, phylogenetic diversity--corresponding to the evolutionary distance between members of a community--offers a useful proxy. Closely related species tend to share traits and interact with their environment in similar ways, but such similarities are lost as evolution progresses. Subtly then, FAB was designed so that bicultures--all plots with just two species--varied widely in their functional and phylogenetic diversity. Some two-species pairs, like white oak and bur oak, were both closely related and quite similar in their leaf shape, nutrient consumption, and responses to environmental stresses like drought and shade. Other pairs of relatively closely related species, like red maple and basswood, differed quite a bit in these traits. Yet other pairs, like basswood and eastern red cedar were both distantly related--remember that the split between flowering angiosperms like basswood and nonflowering gymnosperms like pine took place roughly three hundred million years ago--and functionally distinct. And finally, some pairs of distantly related species, like red oak and white pine, had relatively similar functional traits despite considerable evolutionary divergence. The FAB experiment was designed so that the researchers could determine the role of functional and phylogenetic diversity in bolstering overall productivity. Here, the two-year-old plot in the foreground comprises red oak (Quercus rubra) and white pine (Pinus strobus). 10 Arnoldia 77\/1 ? August 2019 The researchers wanted to understand the impact of biodiversity on herbivore vulnerability and disease susceptibility. Shown here (clockwise from upper left): red maple (Acer rubrum) leaves that have been spotted with anthracnose, an insect gall on a northern pin oak (Quercus ellipsoidalis), and insect herbivory on paper birch (Betula papyrifera), which Grossman evaluates with a plastic grid. Biodiversity Research 11 (This does happen from time to time: consider the functional similarities of bats and insecteating birds.) The presence of these four types of bicultures in FAB allowed us to tease apart the role of functional and phylogenetic diversity in bolstering the ecological functionality of our newly planted \"forest.\" We also wanted to understand whether diversity within a single species--genetic diversity-- was as important as diversity among species. Though intraspecific diversity in other species is often invisible to humans, it has been well-documented that some plant traits vary just as much within a species as among related species. And copious evidence from epidemiology to conservation biology has shown that genetically diverse populations are more stable and better poised to cope with environmental stressors than homogenous ones. To assess this question, we designed and planted a second tree-diversity experiment. The eighthundred-tree Biodiversity in Willows and Poplars (BiWaP) experiment included plots varying not just in species richness but also in genetic diversity. We took advantage of the fact that many species in the willow family (Salicaceae) can be easily propagated by cuttings to grow hundreds of identical clones of several quaking aspens (Populus tremuloides), white aspens (P. alba), and black willows (Salix nigra). We then planted these trees in the field such that some had as neighbors only genetic clones of themselves while others had as neighbors multiple genotypes each of several species. As such, the genetic diversity comprised another dimension of biodiversity whose role in supporting ecosystem function we planned to test. The Complex Role of Biodiversity But what goes into measuring the functionality of an ecosystem--even a highly simplified and orderly biodiversity experiment? At the end of each summer at Cedar Creek, a team of interns--led by me for the first several years of the experiment--measured the stem diameter and height of each tree in FAB. Standardized equations then allowed for easy conversions of these measurements into estimates of trunk biomass. Encouragingly, trunk growth from year to year was higher for trees with morediverse neighbors compared to those in mono- culture (Grossman et al., 2017), although we did not see an effect of either species or genetic diversity in the BiWaP experiment (Grossman and Cavender-Bares, 2019). Paralleling findings from Big Biodiversity, Ecotron, and other grassland experiments around the world, our documentation of a productivity boost in morediverse plots contributed to the growing consensus that this BEF phenomenon is not only confined to grasslands. Indeed, meta-analysis of tree growth data both from global forests (Liang et al., 2016) and managed or experimental systems (Zhang et al., 2012) corroborates our findings. This pattern is perhaps of special note given that monocultural plantations dominate production forestry the world over. Polycultures are harder to maintain and harvest; yet recent experimental findings like ours raise the question of whether increases in yield might compensate for higher costs of maintenance and harvesting. Going beyond my initial focus on productivity, I wanted to determine how tree biodiversity in these systems related to herbivore vulnerability and disease susceptibility. Since we planted FAB inside a massive fenced enclosure, I knew I would never be able to study, for instance, the role of diversity in preventing deer browsing. But I could measure damage by insects and fungal pathogens, like red maple anthracnose. Over three years, I spent a month each autumn painstakingly measuring leaves of hundreds of plants with a translucent grid: I would estimate the original size of a given leaf and the amount of this tissue that had been chewed up by insects or infected by fungi. I also counted galls (small tumors formed by insect larvae) and leaf mines (burrows in leaves created by other larval feeders). Finally, I surveyed damage across the experiment stemming from two fungal diseases, each specialized to a single species in the FAB and BiWaP experiments. The story that emerged from these measurements is a complicated one (Grossman and Cavender-Bares, 2019; Grossman et al., 2018). Having diverse neighbors frequently affected how vulnerable a given tree was to insect or disease damage, but the direction and strength of this relationship varied based on the species of tree and type of damage in question. For instance, having diverse neighbors reduced the 12 Arnoldia 77\/1 ? August 2019 Two genetically distinct white aspen (Populus alba) are shown in the Biodiversity in Willows and Poplars (BiWaP) experiment in the fall of 2016. These aspen look alike when green, but one has attained fall color before the other. likelihood that an oak would be attacked by leaf miners but increased the risk of leaf miner attack for birches! And yes, red maples with more conspecific neighbors were more likely to experience intense anthracnose infection. Fascinatingly, it also seemed that very nearby neighbors (within a one-meter radius of a focal tree) had a bigger impact on that tree's risk of pest attack or disease than did farther away neighbors. This spatial scale-dependence of vulnerability to damage was relatively consistent across tree and pest or disease identity. Generally, though, it appears that other factors, like climate, the presence of predators, and chance, might play a role equivalent to or greater than that of diversity in affecting the vulnerability of trees to pests and pathogens. While pests and diseases constitute the most famous consumers of living plant tissue, an entire food chain unfolds once leaves and roots are shed and begin to decompose, and I also wanted to know how tree diversity affected this microbial universe. Focusing on the rich, plantdependent microbial life of rotting leaves and the soil below them, I was interested in using the FAB experiment as a platform to assess whether more-diverse tree communities might beget more active and diverse soil microbial communities. In both cases, we found subtle biodiversity effects. We found that the most important factor shaping the microbial community was the proportion of trees in a plot that were gymnosperms (pines and junipers) versus angiosperms (oaks, maples, birch, and basswood). Interestingly, pines, and especially junipers, created a hostile environment for bacteria, perhaps due to antimicrobial properties exuded by these species. Yet, since I collected Biodiversity Research 13 samples after only three years of tree growth, it is important to note that the microbial communities of the FAB experiment have probably not finished responding to the presence of different combinations of tree species. So, this story is only just beginning to unfold. Across all these projects, I was surprised to find that species richness--long the standard metric of biodiversity for biologists--emerged as a still-critical predictor of ecosystem function. In study after study, the number of tree species in a plot predicted ecosystem function as well as or better than more abstruse dimensions of biodiversity. In some cases, the diversity of particular functional traits within plots emerged as an important predictor of particular functions. But, generally speaking, I saw little evidence that continuing to measure diversity in terms of species richness might obscure important connections between biodiversity and ecosystem function. From Local to Global to Local Encouragingly, my findings--or anyone else's-- from the tree-diversity experiments at Cedar Creek don't have to be the final word on BEF relationships in forests. On establishment, FAB was inducted into TreeDivNet, a network of twenty-five tree-diversity experiments distributed across the globe. The 1.1 million trees making up TreeDivNet have been planted in sites on six continents and range from boreal to Mediterranean and tropical climates. Though the design of these experiments varies from site to site, each includes some experimental manipulation of tree diversity, as in FAB. At most sites, investigators have made periodic measurements of tree survival and growth, and of damage inflicted upon trees by pests and pathogens (Grossman et al., 2018b). This riot of findings has already contributed to our understanding of how changes in tree biodiversity are likely to affect the way that forests function. And the BEF framework, though developed through experimental work, has now given credence to the idea that biodiversity changes the way ecosystems function. This premise has now been borne out through observational studies of non-experimental (e.g. naturally occurring) ecosystems (van der Plas, 2019). I argue that this holistic view on the value of biodiversity needs to inform the way that we, as managers and users of natural resources, make local decisions. Though large-scale, systemic change will be required for humans to fully address the current biodiversity crisis, such change can be instigated and incubated on the smallest scales. For urbanites, this might mean turning more and more of our marginal spaces into To examine the relationship between tree biodiversity and soil microbes, biodiversity havens. OpporGrossman filled six hundred mesh litterbags with preweighed dried leaves tunities of this nature include varying in diversity that corresponded with the FAB plots. The weight and pollinator-friendly prairie chemical composition of the litter provided a measurement of how decomposition had progressed over the course of four seasons. JACOB MILLER, CEDAR CREEK ECOSYSTEM SCIENCE RESERVE (CC BY-SA 4.0) 14 Arnoldia 77\/1 ? August 2019 Based on the initial results of the FAB experiment, Grossman argues that we should work as individuals and communities to promote biodiversity--not simply within designated natural areas but in yards, parks, and through consumer decisions. yards, urban gardens and food forests, and even no-mow zones such as those currently being put into place at the Arnold Arboretum. Communities can also make choices in our roles as consumers, advocating for less chemically intensive agriculture that protects the incidental biodiversity concomitant with farming prior to the widespread adoption of blanket glyphosate-spraying on row crops. For me, working mere meters away from the Big Biodiversity plots and playing my own part in the establishment of new biodiversity experiments has also highlighted the importance of humility. Empirical evidence shows us that biodiversity plays critical, complex roles in mediating the way ecosystems function. Yet we are often not nor, I would argue, will we ever be able to fully understand and thus manage these BEF dynamics. Instead of assuming that we can figure out how to optimize global biodiversity to provide for the ecosystem functions that we want, it might make more sense to take a precautionary approach. In doing so, we should be highly conservative in both senses of the word, protecting biodiversity far more stringently than we think is necessary to sustain critical ecological functioning, especially in the face of ongoing challenges such as climate change. We would be foolish, I believe, to fail to conserve global biodiversity, which BEF research has shown us to be valuable beyond measure. Acknowledgements The author wishes to acknowledge his doctoral advisor, Jeannine Cavender-Bares, as well as the other co-PIs of the Forests and Biodiversity (FAB) experiment: Sarah Hobbie, Rebecca Montgomery, and Peter Reich. Work described here also stems from collaborations with Peter Kennedy, Jess Gutknecht, and several undergraduate collaborators and interns. Susan Barrott offered helpful feedback on an early draft of this article. Finally, the author is incredibly grateful for the opportunity to work at the Cedar Creek LTER site and to collaborate with TreeDivNet partners. Biodiversity Research 15 References Butchart, S.H.M., M. Walpole, B. Collen, A. van Strien, J.P.W. Scharlemann,... R. Watson. 2010. Global biodiversity: Indicators of recent declines. Science, 328: 1164?1168. Connell, J.H., and E. Orias. 1964. The ecological regulation of species diversity. The American Naturalist, 98: 399?414. Darwin, C. 1859. On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. London: John Murray. Davis, M., S. Faurby, and J. Svenning. 2018. Mammal diversity will take millions of years to recover from the current biodiversity crisis. Proceedings of the National Academy of Sciences, 115: 11262?11267. Downing, J.A., and D. Tilman. 1994. Biodiversity and stability in grasslands. Nature, 367: 363?365. Elton, C.S. 1958. The ecology of invasions by animals and plants. London: Methuen and Co. Fargione, J., D. Tilman, R. Dybzinski, J.H.R. Lambers, C. Clark,... M. Loreau. 2007. From selection to complementarity: shifts in the causes of biodiversity-productivity relationships in a long-term biodiversity experiment. Proceedings of the Royal Society, B 274: 871?6. Grossman, J.J., and J. Cavender-Bares. 2019. Consequences of biodiversity shift across phylogenetic scales for aspen and willow growth, survival, and herbivory. Journal of Vegetation Science, 30: 301?311 Grossman, J.J., J. Cavender-Bares, S.E. Hobbie, P.B. Reich, and R.A. Montgomery. 2017. Species richness and traits predict overyielding in stem growth in an early-successional tree diversity experiment. Ecology, 98: 2601?2614. Grossman, J.J., J. Cavender-Bares, P.B. Reich, R.A. Montgomer y, and S.E. Hobbie. 2018a. Neighborhood diversity simultaneously increased and decreased susceptibility to contrasting herbivores in an early stage forest diversity experiment. Journal of Ecology, 107: 1492?1505. Grossman, J.J., M. Vanhellemont, N. Barsoum, J. Bauhus, H. Bruelheide,... K. Verheyen. 2018b. Synthesis and future research directions linking tree diversity to growth, survival, and damage in a global network of tree diversity experiments. Environmental and Experimental Botany, 152: 68?89. Isbell, F., D. Tilman, S. Polasky, S. Binder, and P. Hawthorne. 2013. Low biodiversity state persists two decades after cessation of nutrient enrichment. Ecology Letters: 454?460. Liang, J., T.W. Crowther, N. Picard, S. Wiser, M. Zhou,... P.B. Reich. 2016. Positive biodiversityproductivity relationship predominant in global forests. Science, 354: 196. May, R.M. 1973. Qualitative stability in model ecosystems. Ecology, 54: 638?641. Naeem, S., L.J. Thompson, S.P. Lawler, J.H. Lawton, and R.M. Woodfin. 1994. Declining biodiversity can alter the performance of ecosystems. Nature, 368: 734?737. Naidoo, R., and W.L. Adamowicz. 2006. Modeling opportunity costs of conservation in transitional landscapes. Conservation Biology, 20: 490?500. van der Plas, F. 2019. Biodiversity and ecosystem functioning in naturally assembled communities. Biological Reviews, brv.12499. Schulze, E.-D., and H. Mooney. 1994. Biodiversity and ecosystem function. Berlin: Springer-Verlag. Tilman, D., F. Isbell, and J.M. Cowles. 2014. Biodiversity and ecosystem functioning. Annual Review of Ecology and Systematics, 45: 471?493. Tilman, D., J. Knops, D. Wedin, P. Reich, M. Ritchie, and E. Siemann. 1997. The influence of functional diversity and composition on ecosystem processes. Science, 277: 1300?1302. Tilman, D., D. Wedin, and J.M.H. Knops. 1996. Productivity and sustainability influenced by biodiversity in grassland ecosystems. Nature, 379: 718?720. Zhang, C., J.A. Postma, L.M. York, and J.P. Lynch. 2014. Root foraging elicits niche complementaritydependent yield advantage in the ancient \"three sisters\" (maize\/bean\/squash) polyculture. Annals of Botany, 114: 1719?1733. Zhang, F., and L. Li. 2003. Using competitive and facilitative interactions in intercropping systems enhances crop productivity and nutrient-use efficiency. Plant and Soil, 248: 305?312. Zhang, Y., H.Y.H. Chen, and P.B. Reich. 2012. Forest productivity increases with evenness, species richness and trait variation: A global metaanalysis. Journal of Ecology, 100: 742?749. Jake J. Grossman is a Putnam Fellow at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"A Medlar by Any Other Name","article_sequence":2,"start_page":16,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25665","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070a76d.jpg","volume":77,"issue_number":1,"year":2019,"series":null,"season":null,"authors":"Enzenbacher, Tiffany","article_content":"ENZENBACHER, T. 2019. A MEDLAR BY ANY OTHER NAME. ARNOLDIA, 77(1): 16?25 A Medlar by Any Other Name Tiffany Enzenbacher T his is an account of a meddling medlar that placed an inquisitive thorn in the side of Jane Ellenbogen Stern, an environmentalist from Pine Bluff, Arkansas. In 1969, Stern was leading a bird-monitoring project at a small remnant of tallgrass prairie and bottomland woods about fifty miles northeast of her home--a natural area now preserved within an expanse of rice fields and aquaculture ponds. She noticed an unusual plant that resembled a hawthorn (Crataegus), only shrubbier, which was covered in white flowers. She collected a branch sample and notified regional biologists, including Edwin Burnell Smith, the curator of the University of Arkansas Herbarium. Stern's discovery triggered the interest of an entourage of plant professionals, who spent nearly a half century attempting to provide a proper identification and name. The plant is now recognized as an unusual naturally occurring hybrid and is known as Stern's medlar (?Crataemespilus canescens). In October 2018, Plant Growth Facilities Manager Kea Woodruff and I eagerly followed Stern's trail to gather propagules from her rare find, during our Arnold Arboretum plant collecting expedition to Arkansas and Oklahoma, which we nicknamed the A-OK expedition. The expedition was part of the Arboretum's Campaign for the Living Collections, a tenyear initiative to collect nearly four hundred taxa from around the globe. We reached out to staff at the Arkansas Natural Heritage Commission in the summer of 2018 to prepare for our upcoming expedition. Theo Witsell, a botanist and ecologist at the commission, explained that Stern's medlar is protected, and as such, we would not be permitted to harvest propagules from the only known wild population growing at the Konecny Grove Natural Area, located in Prairie County. Witsell, however, connected us to his friend Tom Frothingham, a former commission staff member, now of the Little Rock Zoo. Frothingham had obtained a Stern's medlar from the Natural Resources Conservation Service's Plant Materials Center in Booneville, Arkansas, a decade prior. That plant, in turn, had been propagated from the Konecny Grove. Frothingham wrote to Woodruff that, in addition to having a large specimen growing in his yard, he had already potted up a division. \"You'd be welcome to it,\" he wrote, and he also volunteered that we could dig additional divisions. Woodruff and I planned to be near Little Rock during the initial two days of our nine-day expedition, so we arranged to visit Frothingham directly after our six-hour flight (with layover) on October 1. We obtained our rental car and drove forty-five minutes in the unanticipated afternoon heat, starting in the bustling city of Little Rock and continuing through the serene and rural landscapes northwest of the city. We then zeroed in on his address and proceeded to the end of the wooded side road. \"Mine is the second driveway, with the mailboxes,\" Frothingham had instructed. We found our spot. An Unexpected Discovery Born in Little Rock, in 1918, Jane Ellenbogen Stern moved to the suburb of Pine Bluff at about age thirty, with her husband, Howard Stern, and their two children, Arthur and Ellen. Stern developed a long-standing love of the outdoors through birdwatching, which she became passionate about while Arthur was obtaining his Boy Scout nature badge. As her hobby advanced, she became a charter member of the Jefferson County Audubon Society. Coincidentally, Stern was searching for a small bird, the Traill's flycatcher (Empidonax traillii), when she first encountered the medlar. The Traill's flycatcher is now more commonly known as the willow flycatcher. It had been previously documented in the low, moist tallgrass prairies of eastern Arkansas, and Stern was directed to PHOTOS FROM UNIVERSITY OF CENTRAL ARKANSAS ARCHIVES, TORREYSON LIBRARY Stern's Medlar 17 In 1969, Jane Ellenbogen Stern observed an enigmatic shrub in an Arkansas woodland where she was searching for the Traill's flycatcher (Empidonax traillii). Her curiosity about the shrub aroused longstanding taxonomic research. The shrub is now known as Stern's medlar (?Crataemespilus canescens). the Konecny Prairie and Grove--then unnamed and unprotected--which was one of the few remnant prairies remaining in the region. In 1951, a nesting study had documented flycatchers at the site and found that out of the fifteen nests that were discovered, thirteen were in presumed hawthorn trees. One can speculate that the \"hawthorns\" may have been an initial sighting of Stern's medlar. In the summer of 1968, Stern first visited the property, searching for the flycatcher with Raymond McMaster, the manager of the White River National Wildlife Refuge, and Thomas Foti, who would later be appointed senior ecologist for the Arkansas Natural Heritage Commission. Sam Konecny, the owner of the property, showed them around. \"We piled into Mr. Konecny's new car and roared off, helter skelter, across the farm, stopping occasionally to leap out and pull a `weed' while battling off mosquitos and sweltering in the heat,\" Foti later wrote in the Ozark Society Bulletin. The remnant landscape was divided into two sections-- a seventy-one-acre rectangle of tallgrass prairie, which had been preserved as a hayfield (Konecny Prairie), and a twenty-twoacre grove of swampy woods (Konecny Grove), known as slash timber, which included overstory species like persimmon (Diospyros virginiana), green ash (Fraxinus pennsylvanica), and honey locust (Gleditsia triacanthos). Along the edge of this woodland were the hawthorn-like thickets in which the flycatchers nested. The following winter, Stern was contacted by Douglas James, a professor of zoology at the University of Arkansas, who asked her to organize an effort to determine when the Traill's flycatcher arrived at the grove. The reason for the effort was due to uncertainty about flycatcher taxonomy. Ornithologists increasingly believed the Traill's flycatcher should be treated as two species, rather than one, but they were uncertain which species deserved the original scientific name, which was based on a bird John James Audubon had observed in Arkansas. The song is considered the best way of distinguishing between the two species. The population that was already known to breed in Arkansas sang \"fitz-bew,\" but the popula- UNIVERSITY OF CENTRAL ARKANSAS ARCHIVES, TORREYSON LIBRARY 18 Arnoldia 77\/1 ? August 2019 Stern organized a group of volunteer bird-watchers in the spring of 1969. The volunteers visited the Konecny Prairie and Grove Natural Area--then unofficially named--to monitor the arrival of the Traill's flycatcher. tion that sang \"wee-be-o\" had previously not been reported to nest in the state. Due to Stern's experience and proximity to the site, she was the ideal individual to lead the effort. She was eager for the challenge. \"We will do our best to cover the Konecny place,\" she wrote to James on February 26, 1969. \"If the fiz-bew one hollers as loud as the spit-chee and che-bek and wee-bee-o, I don't see how anyone could miss him.\" Several months later, the \"Traill's Flycatcher Vigil\" commenced. Stern assembled twentyseven birdwatchers to take turns observing the grove daily, beginning March 29, to determine when the flycatchers arrived to roost. On May 6, Stern was on the lookout with Jewel Herring, another birder from Pine Bluff, when they heard one of the flycatchers singing, and the song was \"fitz-bew.\" On July 7, Foti and Stern returned and found an empty nest, assumed to UNIVERSITY OF CENTRAL ARKANSAS ARCHIVES, TORREYSON LIBRARY Stern's Medlar 19 \"Originally he had it perched on a honey locust twig,\" zoologist Douglas James wrote to Stern, describing an illustration of the Traill's flycatcher. \"I returned it saying good grief it's the hawthorn that determines the bird's presence on the Grand Prairie.\" James, however, was uncertain whether the illustration matched the unusual hawthorn-like shrub Stern had observed, and he requested her confirmation. be a Traill's flycatcher's, burrowed in a branch of a hawthorn-like shrub. Stern observed that the plant did not appear like other hawthorn species in the grove, and so she sent a sample to Edwin Burnell Smith, the curator of the University of Arkansas Herbarium. Stern would ignite a decades long process of pursuing an accurate name for the plant, even while the confusion around the flycatcher taxonomy was resolved. The \"wee-be-o\" species, commonly known as the alder flycatcher, was given the official name of Empidonax alnorum in 1973. The Search for a Name On July 10, 1969, Smith wrote to Stern with an initial classification of the mystery plant. \"I must say, [this is one] of the most difficult plants I've received for identification,\" Smith wrote. \"The small tree is a type of Crataegus, `Hawthorn,' in the Rose Family. The genus Crataegus is cursed with a very complex taxonomy which makes the individual species quite difficult to identify. The one you sent in is probably (and I emphasize probably) Crataegus engelmannii.\" As Smith was unsure of his initial naming, he solicited an additional sample and offered another stab--hillside hawthorn (C. collina). Smith even mailed an herbarium sheet to the Arnold Arboretum for assistance, likely due to the reputation of Charles Sprague Sargent, the Arboretum's founding director, as an expert on hawthorn taxonomy. Director Richard Howard responded to Smith on October 23 with a complicated assessment: it appeared to be a \"mixed collection.\" The flowering stems looked like red chokeberry (Aronia arbutifolia), and Howard agreed that the fruiting stems (obtained that fall) resembled C. collina. \"Unhappily,\" 20 Arnoldia 77\/1 ? August 2019 Howard wrote, \"the specimens lack the young leaves and flowers which are equally critical for accurate determination.\" Correct identification was proving to be exceedingly difficult, and the only sense that could be made was that Stern was harvesting samples from multiple plants. However, in a letter to Stern on June 8, 1970, Smith suggested an alternate explanation for the \"enigmatic `Haw'\"--the possibility of \"a strange hybrid of some kind.\" But then, after studying additional collections, Smith was relieved to finally provide the name of dotted hawthorn (Crataegus punctata). Dotted hawthorn is very similar to C. collina, and is occasionally considered the same species. \"Well, finally (!) I am able to report to you with great relief that I have been able to determine the frustrating `haw,' at least to my satisfaction,\" Smith wrote to Stern on October 15. \"It turns out that the plants are not a new species or a hybrid, which is kind of sad after all the trouble we have both gone to.\" Later, upon reexamination of specimens for the Vascular Flora of the Southeastern United States in the 1980s, taxonomists rejected Stern's medlar as a hawthorn. Although the flower and fruit characteristics are similar to hawthorns, as is the overall plant height, hawthorn leaves are shallowly to deeply lobed, unlike the simple, subentire leaves of Stern's medlar. Also, hawthorns typically grow as small trees with one large stem, occasionally producing suckers, while Stern's medlar is a large shrub with equal-diameter shoots. Taxonomist James B. Phipps, of the University of Western Ontario, was intrigued by the anomalous specimens, and contacted Smith in 1988. Phipps had been researching hawthorn taxonomy since the 1970s, and years later, he would author the hawthorn section of the Flora of North America. Smith relayed the news to Marie Locke of Pine Bluff in a letter dated July 13, 1988. \"I recently heard some interesting news from a botanist in Canada: He plans to name a new species (and new genus, for Arkansas) in the Rose family from collections made several years ago by Jane Stern,\" Smith wrote. \"I think that this is the plant I had such difficulty identifying-- remember it? It had two strikingly different kinds of leaves on it.\" Stern was later in contact with Phipps herself. She agreed to collect additional herbarium specimens in mid-September, when the fruit would be ripe, and she also promised to arrange for Phipps to visit the Konecny property himself. On October 21, a group of Arkansans, including Stern, escorted Phipps to the coveted shrubs. Phipps ultimately published a new name for the plant in 1990: Mespilus canescens. He recognized that although Stern's plants resembled hawthorns, other characteristics uniquely resembled common medlar (Mespilus germanica), the singular species within that genus, which is native to southeastern Europe and Iran. Among other things, it shared a multistemmed habit; distinctive leaf venation, with secondary veins curving toward the margins; and fine, white (canescent) hairs on the inflorescence. While even this comparison didn't match completely--the common medlar, for instance, produces larger brown fruit, unlike the red fruit of Stern's discovery--Phipps suggested that a hybrid origin seemed unlikely, given the lack of suitable parent species. \"The most likely explanation of the status of M. canescens is that it is an ancient relic,\" Phipps wrote. \"One should always be cautious in describing a new species from such limited material (all the cited collections come from the same locality), but M. canescens is so distinct from all other native American Maloideae that there can be no doubt that it is not a previously described North American member of this subfamily.\" In the same paper, the shrub was aptly given the common name of Stern's medlar. But the story was not over. Almost twenty years later, Eugenia Y.Y. Lo and colleagues further investigated the relationship of Stern's medlar to the common Eurasian medlar. Through DNA amplification and phylogenic analyses of over ninety Rosaceous species, including hawthorn, chokeberry, crabapple (Malus), medlar, and serviceberry (Amelanchier), they concluded that although Stern's medlar shares a common ancestor with the Eurasian medlar, it is more closely related to blueberry hawthorn (Crataegus brachyacantha)--a species whose range is centered in Louisiana, eastern Texas, and southern Arkansas. Their analyses, published in 2007, suggested a hybrid origin of Stern's med- UNIVERSITY OF CENTRAL ARKANSAS ARCHIVES, TORREYSON LIBRARY In 1988, taxonomist James B. Phipps visited Konecny Grove with Stern and other local environmentalists. Archival correspondence revealed the back-and-forth excitement, with a letter from Stern to Phipps (top left), coordinating Phipps's visit, and a letter from Harold Grimmett, the director of the Arkansas Natural Heritage Commission (bottom right), providing a collecting permit for a follow-up visit. Phipps proposed that the plant was a new species of medlar, which he named Mespilus canescens. 22 Arnoldia 77\/1 ? August 2019 tion, and aid from government agencies, private land corporations, and the railroad industry. Perhaps the families in Slovak had brought along plants or seeds of a favorite fruit? Common medlar has been cultivated as far back as the ancient Romans. Phipps accounted for these new conclusions, and in 2017, he reclassified Stern's medlar as ?Crataemespilus canescens. The ?Crataemespilus nothogenus was created in 1899 to accommodate an assumed hybrid, ?C. grandiflora, originating from midland hawthorn (Crataegus laevigata) and common medlar, that was also initially described as medlar. In 1914, a second hybrid was discovered: ?C. gillottii, an intermediate between English hawthorn (Crataegus monogyna) and common medlar. Now Stern's medlar has joined the ranks as the third member of this hybrid genus. WONDERLANE, CC BY 2.0 lar, with blueberry hawthorn as the maternal parent. Because the fruit of blueberry hawthorn is fittingly blue, however, the authors acknowledged that another native red-fruited hawthorn (or even an ancient, now extinct medlar species) may have been involved in past hybridization. The authors surmised that common medlar and blueberry hawthorn may have hybridized if they were cultivated within range of one another. Hawthorns are known to hybridize, and the authors pointed to literature confirming that common medlar was, indeed, cultivated in an agricultural station in Louisiana as far back as 1893. Furthermore, Slovak, the small town two miles north of Konecny Prairie and Grove, was home to at least fifty families of Eastern European heritage by 1909. Immigration to Arkansas in the mid- to late-nineteenth century was encouraged by advertising, legisla- Common medlar (Mespilus germanica) is harvested while unripe and allowed to blet (soften). After storing in a cool dark place until squishy and aromatic, the fruits are ready for direct consumption or for use in jellies or wine. LARRY ALLAIN, U.S. GEOLOGICAL SURVEY TIFFANY ENZENBACHER Stern's Medlar 23 Through genetic analyses, researchers have now determined that Stern's medlar (?Crataemespilus canescens, right) is a hybrid between the common medlar and the blueberry hawthorn (Crataegus brachyacantha). An Unconventional Collection The taxonomic and conservation status of the Stern's medlar initially placed this plant on the radar of the A-OK expedition. When Woodruff and I pulled into the driveway at Tom Frothingham's property, northwest of Little Rock, we were greeted by him and two colleagues, Lauren Goldstein and Connor Livingston. Woodruff and I could hardly contain ourselves as Frothingham led us from the driveway to where the specimen was planted, out in full sun, between the shed and house. The shrub was vigorous--it was nearly fifteen feet tall--and I thought its habit resembled that of a serviceberry or large rose (Rosa)--upright with slightly cascading branches. After months preparing for the A-OK expedition and a day's worth of travel, it was surreal that our target was in plain sight. To add to our excitement, the medlar was fruiting! Frothingham and I alternated ascending the ladder to gather the cherry-sized pomes, which were relatively sparse. After we collected a handful, Frothingham insisted that the whole group sample one, to which we all curiously obliged. The shiny red fruit was surprisingly sweet, and Woodruff and I saved all the seed from the consumed fruit to send back to the Arboretum's Dana Greenhouse for propaga- tion. I then slowly walked around the specimen and found an appropriate division to dig. Frothingham lent me a trowel for the job. After the division was successfully dug and bagged, Frothingham then led us to the promised potted plant, harvested as a division several years prior, at the front of his house. We chatted about seed propagation and the rich history of the plant for a while, before Woodruff and I loaded up our bounty. We repeatedly thanked our collaborator for his generosity, and we backed out of his driveway, delighted about how successful the first day of the expedition had been. Preserving an Unusual Hybrid Not only is Stern's medlar a rare hybrid but it is visually appealing to boot. In September 1989, Stern wrote to Harold Grimmett, then the director of the Arkansas Natural Heritage Commission, urging him to request that Phipps withhold the location of the Stern's medlar in his 1990 paper. \"The plant is extremely attractive in appearance and can be expected to be aggressively sought by the horticultural trade,\" Stern wrote. Phipps, in response, suggested that the commercial threat seemed \"highly unlikely\" and noted that the location was already well-documented with herbarium spec- KEA WOODRUFF 24 Arnoldia 77\/1 ? August 2019 flycatcher migrated elsewhere to nest, and the landowner told Stern that he was interested in reclaiming the property. \"So many things connect with the Konecny prairie, the grove, the bird, and the Tree,\" Stern wrote to Phipps on April 24, 1989. \"Bless the bird for hanging on long enough for the Commission to purchase the easement on the grove ... He would have plowed some or all of it ... but you and The Tree have put an end to that idea.\" Stern's medlar is now graded as critically endangered, which means it has an extremely The author harvests a division from the Stern's medlar (?Crataemespilus caneshigh risk of extinction in the cens) at Tom Frothingham's property on the outskirts of Little Rock, Arkansas. wild. Only twenty-four indiThe division was bagged and mailed for next-day delivery to the Arnold Arboreviduals are known to exist, and tum's Dana Greenhouse. because Stern's medlar is tripimens. Nonetheless, Phipps agreed with Stern's loid (having three sets of chromosomes), it is assessment of the plant's beauty. In his book likely sterile. This means the seeds we collected Hawthorns and Medlars, published in 2003, from Frothingham's specimen will be difficult Phipps states that \"Stern's medlar is arguably (if not impossible) to germinate. As hawthorns the most exquisite ornamental treated in this have been found to produce seed through apobook.\" Its exfoliating bark has hues of cream mixis (asexual seed formation), however, hope and olive, and he describes the plants as \"a for potential seedlings is well-founded. Regardfountain of white flowers.\" I personally like to less, the clonal division that we harvested, as imagine that this attractive nature is the reason well as the potted plant that Frothingham prothat Stern initially took such keen interest in vided, are thriving at the Dana Greenhouse prothe plant--launching a pursuit that continued duction facility. In two to three years, they will well beyond the Traill's flycatcher. be added to the Arboretum's living collections In fact, the medlar has proved to be a conto join the other 179 taxa in our landscape that servation boon for Konecny Prairie and Grove. are of conservation concern. I am proud to have From the beginning, Stern recognized the signifbrought Stern's medlar to the Arboretum with icance of this site as the only remaining Traill's Woodruff, to have the opportunity to learn of flycatcher's nesting habitat in eastern Arkansas Stern's tireless conservation efforts, and to have and as one of the few tallgrass prairie remnants experienced, first-hand, what Phipps devoted that escaped plowing in the region. Stern awakhis entire career to--the complicated identificaened a movement to preserve the prairie and tion of hawthorns, medlars, and their hybrids. grove, and in February 1976, the grove became Works Cited the first conservation easement purchased by the Arkansas Natural Heritage Commission. All archival correspondence and images were provided This was a victory for so many, and on May 14, courtesy of the Jane E. Stern Collection, University of 1977, a gathering was held on the Konecny land Central Arkansas Archives (M90-02, Series II, Sub-Series to celebrate, which coincided with the Traill's IV, Box 1, Files 12?17), Torreyson Library, University of Central Arkansas, Conway, Arkansas. flycatcher arrival. A decade later, however, the TIFFANY ENZENBACHER Stern's Medlar 25 Tom Frothingham (at left) encouraged the collectors to sample fruit of the Stern's medlar (?Crataemespilus canescens). His colleagues Lauren Goldstein and Connor Livingston are pictured, along with Kea Woodruff (right). Foti, T. 1971. The grand prairie. Ozark Society Bulletin, 5(4): 6?11. Freeman, F.D. 1948. Immigration to Arkansas. The Arkansas Historical Quarterly, 7(3): 201?220. Kesterson, K. 2019. Jane Rita Ellenbogen Stern (1918? 1989). In CALS Encyclopedia of Arkansas. Retrieved from https:\/\/encyclopediaofarkansas. net\/entries\/jane-rita-ellenbogen-stern-421\/ Joshi, V.K., and Attri, B.L. 2017. Specific features of table wine production technology. In Kosseva, M.R., Joshi, V.K., and Panesar, P.S. (Eds.), Science and Technology of Fruit Wine Production (pp. 295? 461). Cambridge, MA: Academic Press. Leslie, J.W. 2000. Stern, Jane Rita Ellenbogen. In Williams, N.A., and Whayne, J.M. (Eds.), Arkansas Biography: A Collection of Notable Lives (pp. 274? 275). Fayetteville: University of Arkansas Press. Lo, E.Y.Y., Stefanovic, S., and Dickson, T.A. 2007. Molecular reappraisal of relationships between Crataegus and Mespilus (Rosaceae, Pyreae)-- two genera or one? Systematic Botany, 32(3): 596?616. Meanley, B. 1952. Notes on nesting Traill's flycatcher in eastern Arkansas. The Wilson Bulletin, 64(2): 111?112. Phipps, J.B. 1990. Mespilus canescens, a new Rosaceous endemic from Arkansas. Systematic Botany, 15(1): 26?32. Phipps, J.B. 2016. Studies in Mespilus, Crataegus, and ?Crateamespilus (Roseaceae), I. Differentiation of Mespilus and Crataegus, expansion of ? Crataemespilus, with supplementary observations on differences between the Crataegus and Amelanchier clades. Phylotaxa, 257(3): 201?229. Phipps, J.B., Kennon, N., and Lance, R. 2003. Hawthorns and medlars. Portland: Timber Press. Vaughn, M. 2015. Hawthorn: The tree that has nourished, healed, and inspired through the ages. New Haven: Yale University Press. Tiffany Enzenbacher is the manager of plant production at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"A Botanist in Borneo: Understanding Patterns in the Forested Landscape","article_sequence":3,"start_page":26,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25664","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070a728.jpg","volume":77,"issue_number":1,"year":2019,"series":null,"season":null,"authors":"Ashton, Peter Shaw","article_content":"ASHTON, P. 2019. A BOTANIST IN BORNEO: UNDERSTANDING PATTERNS IN THE FORESTED LANDSCAPE. ARNOLDIA, 77(1): 26?39 A Botanist in Borneo: Understanding Patterns in the Forested Landscape Peter Ashton W scapes of mild, moist climates are universal. Yet, in Brunei, I began to sense that individual species within these rainforest communities were often more highly habitat specific than I had ever seen in temperate forests. After twenty-eight intensive months of fieldwork, camping, and longhouse life in Brunei, I made the case that differentiating between these inland forest types--known collectively as mixed dipterocarp forests (or MDFs)--could have important implications for timber inventories and silviculture. At that time, timber was only cut for local use in northwestern Borneo, although research towards sustainable harvesting was advanced in Peninsular Malaysia. Even there, as elsewhere in the tropics, distinct types of lowland MDFs had yet to be defined. The forestry department gave me clearance and funding to lay out sets of plots to test my hypothesis. For foresters, understanding the distribution of these tree communities could guide sustainable harvesting practices. But knowledge of tree species preferences and distributions would also provide the means of mapping biodiversity, locating centers of richness and endemism, and identifying and demarcating priorities for conservation--a first for the tropics. hen I began as a field researcher on the island of Borneo, in 1957, little was known about the distribution of the inland rainforests. The forests are incredibly diverse and are dominated by large overstory trees in the dipterocarp family (Dipterocarpaceae), which often tower more than two hundred feet above the forest floor. While variation in the forests was evident, it appeared as chaotic and random as the colored specks in a children's kaleidoscope. Odoardo Beccari, a Florentine botanist who spent two years in northern Borneo in the 1860s, had provided the first confirmation that specialized lowland habitats, including peat swamps and sandy exposures, bear distinct forest types, but neither he nor his successors until my time had recognized any correlation between habitat characteristics and forest structure on the yellow-to-red tropical soils that characterize much of the inlands. Yet as I tramped along Bukit Biang--a long ridge in eastern Brunei--I was surprised to see dipterocarp species I had come to know on the sandy coastal hills of western Brunei, and I began to sense that these forests were divided into two distinct communities--one on sandy soils, the other on loams. These communities would reappear in different localities as I extended my explorations throughout Brunei. I came to anticipate the flora by the distinctive sounds of the cicadas that inhabited each and by the smells of the forest, which I later recognized all over Borneo and even Peninsular Malaysia-- the mellow fruitfulness and fermentation from the loams or the resinous aroma from the peaty humus covering the sandy soils. Those forest smells returned to me decades later, after I had assumed the directorship of the Arnold Arboretum and first trekked into the loamy bottomlands in the Connecticut River Valley and the sandy pine barrens of New Jersey and Cape Cod. It was only then that I came to understand that these habitat patterns in the forested land- Work on the research plots commenced in 1959. But first, I briefly returned to England for my own wedding. My wife, Mary, was to become the perfect companion for a life of jungle exploration. She had been born and spent her first years in Sri Lanka, where her family had been in trade and tea for over a century. She would join my hectic field life at once, and we only had a few days in town before departing for a long stay in the hulu (or upriver country). I wanted to document and compare the two main forms of MDFs that I had recognized in my explorations. I decided to compare two seemingly contrasting sites--one on the sandy coastal hills of Andulau, in western Brunei, and the other on 50 miles 100 kilometers Bandar Seri Bagawa Bagawan awan wan n BRUNEI Andulau An nd n d dul u Forest orre re t Reserve R eser serrve Miri Kual Kuala K Ku ual ala laa Be B Belalo eel ela llalong lal la alo on ng ng Lambir Lam L La am mbiir Hills Hi lss H SOUTH CHINA SEA Ulu Ul U lu Ba Bak Bakong akon ak ng ng Bok-Tisam Bo ok Ti Ti am m Fo Forest ores e R Reserve eser ese erv Iju ju Hill Hiill H Moun Mount Mo M oun o nt Me nt M Mersing rsing rsin si g si Sibu ibu ibu ib u Raja ng River Santubongg Mountain t Bara er Riv Sega Segan S Seg ggaan a F Forest res ess Res ve Reserve Res r ve Nyab Nyabau Nya N Ny yyabau ya baau uF Forest ore rees est st Re Res Reserv Reserve es rrvee m Tin jar Ri Ulu Ul U lu D Dapo Dapoi po p oi oi MAL AYSI A Sarawak Cara Carapa C Car arrap aapa p paa Pi P Pila iila l la Raya R Ra ayya Hil aya Hill Hi H ill lll Ulu U Ul lu M Mujo Mujong ujjong n ngg Bako National Park Kuch Kuc Kuching K Ku ucchiiingg u L u p a r R i ve r BORNEO I N D O NE SI A Peter Ashton's research identified distribution patterns in the hyperdiverse inland rainforests of northern Borneo, collectively known as mixed dipterocarp forests (MDFs). His research ranged between sites like the Andulau hills, in western Brunei, where forests occur on sandy soils (above left), and Carapa Pila, in central Sarawak, which supports large trees like Shorea mujongensis (above right) on loam soils. Ashton's research sites are mapped, along with his primary expedition routes (green) and his shorter secondary routes (blue) between 1963 and 1966. MAP BY PETER ASHTON, ARNOLD ARBORETUM, AND GIS COMMUNITY ALL PHOTOS BY AUTHOR FROM ARNOLD ARBORETUM ARCHIVES Peter Ashton in Northwest Borneo 27 28 Arnoldia 77\/1 ? August 2019 After their marriage in 1959, Mary Ashton joined Peter in the field. Here, Mary is shown with longtime field assistant Asah anak Unyong at Kuala Belalong, a research site on loam soils in eastern Brunei. Asah collects Borneo fiddleheads (Diplazium esculentum) for supper at the same site (below), while another collector poses a flowering branch of Dillenia excelsa (Dilleniaceae), a large tree observed on the sandier soils of the Andulau hills. Peter Ashton in Northwest Borneo 29 steep clay-loam ridges near Kuala Belalong, in eastern Brunei. I sensed that topography was an important feature, causing local variation in the form, composition, and possibly growth rates of the forests, so at each site, I planned for fifty one-acre plots that covered the complete topographic spectrum (including ridgetops, slopes, valley bottoms, and riverbanks). Each one-acre plot represented a homogeneous topography, and the number and size of the plots were visually estimated to be representative of the forest variation at each site. This method had already become a standard procedure for ecologists studying temperate grasslands, but it was a first for biodiverse tropical forests and, indeed, for any forests known to me. All trees above one-foot circumference were to be documented. We measured the trunk girth and estimated the height for each. We also identified each as morphologically different species by means of fallen leaves and local Iban names. But I rarely knew the scientific name for these visually distinct trees. Life eventually settled down to calm and peace at out first location, Kuala Belalong. It was one of the most beautiful places in which we ever camped, with the rush of water over the rapids in our ears as we lived and slept. While I scrambled daily up the muddy slopes, locating plot positions, surveying, and initiating documentation with the teams, Mary would take her typewriter to the shingle beach, preparing herbarium labels and editing field notes. She saw the wildlife that I rarely or never experienced: a pair of small-clawed otters (Aonyx cinerea) that came to join her, squeaking and gamboling in the shallow water nearby; the extraordinary and terrifying pack of Bornean beared pigs (Sus barbatus), thundering headlong down the steep hillside opposite, then splashing across a rapid upriver, on migration in search of fruiting trees; and a macaque (Macaca fascicularis) who discovered our food store and made off with some delicacies. Our work in Andulau began the following year, in 1960, and was carried out smoothly, given our growing experience and the gentle landscape. I continued to conduct multi-week collecting expeditions until I was satisfied that the diversity of landscapes and forests in Brunei had been examined. When we returned to the University of Cambridge, after the 1960 field season, I brought back two suitcases of fallen leaves and twenty notebooks--all needed to complete my doctoral dissertation. I had presorted the leaves into nearly eight hundred morphospecies, which were recognizable entities that were mostly (dipterocarps excepted) lacking a formal scientific name and therefore named using indigenous nomenclature. They were to be named with the help of taxonomists at the University of Leiden and the Royal Botanic Gardens, Kew, supported wherever possible by our herbarium-quality specimens in flower or fruit. This groundwork provided the basis for a Checklist of Brunei Trees, which I co-authored with Hasan bin Pukol, the ascendant curator of the Brunei Herbarium, who had been a mentor on tree identification from the time he joined our team. He was an experienced informant on traditional plant uses and became a good friend who involved Mary and me in his family activities, including marriages and births (and he even got us invited to a royal circumcision). Our combined efforts eventually provided accurate names to support the baseline data from our one hundred plots: some twenty-five thousand individuals representing nearly seven hundred species. I had no idea how I could sort and compare the plots using this elephantine data set! But luck came my way, for a few months after my arrival in Cambridge, the annual meeting of the British Ecological Society was convened there. I attended, where I was recommended to confess my rash achievements to Peter Greig-Smith, a professor at the University College of North Wales, Bangor, who was one of the leaders of a new science known as quantitative plant community ecology. Greig-Smith, a modest if somewhat austere academic, asked whether I had seen the recent paper, published in Ecological Monographs, on the woodlands bordering the tallgrass prairies of northern Wisconsin. Authors Roger Bray and John Curtis of the University of Wisconsin were to save my career. They had devised a method of relating their forests to one another, and to their climate and soils, by comparing plots according to the presence and abundance of each species ADAPTED FROM ASHTON, 1964 30 Arnoldia 77\/1 ? August 2019 BELALONG 7 HILLSIDES ca. 450 meters 6 Increasing steepness Decreasing humus 5 RIDGES ca. 600 meters HILLSIDES BELALONG RIVER BANK Decreasing steepness RIDGES RIDGES ca. 200 meters ca. 650 meters ca. 450 meters 4 RIDGES TEMBURONG RIVER BANK Increasing humus ca. 550 meters 3 2 3 4 5 Very humid sticky soils, shallow roots, poor drainage pH ca. 4.6 6 7 8 Friable silty soils, deep rooting, excessive drainage pH ca. 4.2 Ashton used a statistical method known as a Bray-Curtis ordination to visualize the relationships between habitat characteristics and the tree communities for two research sites in Brunei. Here, fifty plots from Kuala Belalong have been organized into groups using this method. present. This general method is known as ordination, and elements of their novel approach are still used today. Examination of the data from Brunei indicated that the tree flora at Belalong was markedly different from that at Andulau, with only a third of species common to both sets of plots, so I decided to ordinate the two sets separately. I set to work, entering the plot data and calculating the matrix of similarity indices. I toiled night and day, for more than three weeks, using a bulky hand calculator, the Swedish Facit. The day of reckoning arrived when I started to place the plots, as dots, using a simple geometric technique for ordination, in two dimensions so that the distance between them related to their floristic similarity. To my amazement, a recognizable pattern gradually emerged for the fifty plots at each site, much like, in those days, how a photographic print would emerge on paper set in hypo solution. The patterns confirmed intuitions gained from field experience, with the tree flora tied intimately with geology and topography. Although we hadn't recensused the plots to show change over time, the initial results were clear. The species composition indicated that the dipterocarp species on clay loams, and particularly the lower slopes and undulating land, were predominantly light hardwoods that grow relatively fast. These species might yield a timber crop within a half century. But the sandy soils of Andulau, and also the shallow loams along the sharp Belalong ridges, were dominated by heavy dipterocarps and other hardwoods whose growth rates were known from other research to be much slower, implying that more complex management would be needed to sustain selective felling, with growth cycles exceeding a century. Peter Ashton in Northwest Borneo 31 The Ashtons returned to Borneo in 1962. Here, Mark Ashton, at age four, carries a macaque (Macaca fascicularis) named Mr. Nips. Mark currently professes silviculture at Yale, where he also directs the Yale School Forests. Peter Ashton's research sites ranged from coastal locations like Bako National Park (top) and mountainous inland locations like Carapa Pila (bottom). expanded--both geographically, to see whether the same trends would be observed elsewhere on Borneo, and temporally, to observe changes in the forest structure over time. The ordinations demonstrated, for the first time, that hyperdiverse tropical lowland forests were as floristically variable and habitatspecific as temperate broadleaf forests; indeed, the individual species showed a degree of habitat specificity only found in temperate forests in specialized habitats such as limestone crags. It is the tree species, through their chemical and physical interactions with other forms of life, that directly or indirectly mediate all biodiversity. This knowledge provided a breakthrough, and I knew that the work needed to be Mary and I, now with a young family, returned to Borneo in 1962. I resumed my explorations, this time as forest botanist in the adjacent state of Sarawak (which became part of Malaysia in 1963), where we eventually spent five years. It was a perfect place to bring up our three children. Soon we were taking them to the woods where 32 Arnoldia 77\/1 ? August 2019 we, looking up into the canopy for signs of flowering, would be nudged by them, looking down with a detail to be observed from their stature alone, alerting us to a nest of giant ants, or a huge millipede, or a leech swaying encouragingly. Sarawak is slightly larger than New York State and therefore twenty times the area of Brunei. This presented both a challenge and an opportunity. I adopted a plan whereby, as in Brunei, I would undertake one major botanical exploration each year, while making periodic short forays when time allowed. Although Sarawak and Brunei share a dominant sedimentary geology of sandstones, shales, and clays, Sarawak also supports tantalizing habitats on isolated pockets of limestone karst and volcanic rocks. I further suspected that the major waterways--the Baram, Rajang and Lupar Rivers--could provide evidence of separate diversification, even speciation, in the lowland forests isolated on either side of their extensive floodplains. The major expeditions provided opportunities to set up plot clusters in a diverse set of habitats and geographical locations. As in Brunei, plot results would add to knowledge needed to understand patterns of timber supply and quality. We would also have opportunities to initiate recensusing of permanent plots, conducted on five-year intervals. This would allow us to test predictions of growth rates and management protocols inferred from the static data gathered in Brunei. Our first year of plot surveys started with a crisis: The December-to-February northeast monsoon came in with a fury not matched in recent history. Kuching, in the west, where we all lived, received continuous rain rattling on our roofs for more than two weeks at a time. Over one-third of the state was underwater, but miraculously, no lives were lost thanks to the army with their inflatable rivercraft. But it also provided an opportunity to establish permanent plots on landslide locations, uniquely allowing us to monitor forest regeneration from scratch. Afterwards, these journeys became routine, socially as well as botanically. For the first Upriver travel occasionally proved dangerous and taxing. Here, one of Ashton's boats approaches the head of a rapid, Ulu Mujong, in central Sarawak. Peter Ashton in Northwest Borneo 33 few hours upriver, travelling in a dugout canoe known as a prahu, we would pass through mangrove forests to the first Malay fisherman's town where we would check in at the government office, buy food, and learn whether our advance party had succeeded in attracting local Dayaks--the indigenous people of Borneo--who were familiar with the terrain and who would also provide boats. We would then proceed to the forest, often spending nights in riverside longhouses along the way. Each longhouse is essentially a village of wooden rowhouses, on stilts and under one roof, entered by ascending a notched log (like a ladder) at either end. The shared roof shelters a gallery on whose floor all social activities flourished. These stayovers were always hilarious, if somewhat raucous, experiences. As the chosen house approached, our team of local field assistants would begin combing and oiling their hair, smartening up and adding perfume; when we turned the corner and saw the longhouse for the first time, as often as not there would be a group of young women, having heard the noise of our outboards, who would have descended to the landing to wave and shout encouragement. Conversations on these boat rides were always alive with joking and good-natured braggadocio. At first, the riverside vegetation would consist of a mix of cultivated trees--indigenous mango and durian species, rambutans, coconut palms, and native and Cavendish bananas-- and the indigenous species of the floodplains. But things would change as soon as the current quickened and the first rocky banks were exposed. A distinct flora appeared below the flood line: miniature palms, aroids, ferns, and a diversity of shrubs and coarse herbs, known as rheophytes, which are adapted to periodic immersion and sweeping floodwaters--a community rich in rare species awaiting collection. Overhead, trees that had gained traction on the rocks leaned precariously over the narrowing water, their branches dripping with epiphytic ferns, orchids, and even rhododendrons (Rhododendron sect. Vireya). We Supplies were carried in woven rattan packs known as selabits, shown with field assistants at Ulu Mujong (left). The work resulted in massive amounts of specimens. A field assistant, Brain, arranges and prepares specimens near the coastal town of Bintulu. 34 Arnoldia 77\/1 ? August 2019 Ashton's research ultimately documented around twenty-five hundred tree species at research plots in Sarawak and Brunei. Clockwise from top left: Scyphostegia borneensis (willow family, Salicaceae), Dysoxylum sp. (mahogany family, Melieaceae), Melanorrhoea inappendiculata (cashew family, Anacardiaceae), Ixora sp. (coffee family, Rubiaceae), Didesmandra aspera (Dilleniaceae), and Sterculia megistophylla (mallow family, Malvaceae). Peter Ashton in Northwest Borneo 35 solution. Everything--everything--had to be kept waterproof, in camp and in transit, and this required commissioning special durable waterproof backpacks. This work became my ecological specialty. We ended up with 105 plots, each 1.5 acres (increased from the Brunei experience), at thirteen sites in eleven localities. The work at each site, following establishment, which I personally undertook, was carried out by my team of climbers, some of whom had rejoined me when they heard of my return, and was led by an experienced Sarawak Malay forester. Among these foresters, I most remember Ilias bin Pa'ie, keeper of the Sarawak herbarium, who was a close friend and mentor, ever cautious and gentle, who tragically died from a heart attack when overseeing the 1975 recensusing at Lambir, a hilly site in the northeastern corner of the state. While this was some years after my departure, the loss was profound. observed trees like Dipterocarpus oblongifolius, which produces bright-pink winged fruit that hang like Christmas decorations, and the fragrant babai (Saraca declinata), a leguminous species, which produces decorative yellowishorange flowers on its trunk. At this point in the upriver journey, those with boating skill came to the fore, led by the outboard operator and the individual stationed on the prow with a long suar--a fending pole-- who was known as the jagar luan (or prowguard). The most crucial moment was always at the head of a rapid when the prahu, lifted during its ascent, would drop onto the calm water as we entered a lagoon. The stern would lift, and with it the propeller. All hands took to the poles for, if we failed to heave the baggage-filled prahu across, we could lose control, fall back headlong, probably sideways, and lose everything to the torrent. I experienced this seven times, but thanks to willing and experienced hands, we never lost our precious specimens, notebooks, or soil samples, although a camera could get a dousing. I would watch and admire the skill shown. But upriver travel was punishing to the outboard motor: what would start as a shiny new Evinrude with fancy hood would, on occasion, end up as an unprotected swirling stick, topped with a greasy bareboned engine-- still miraculously spinning! We aimed to establish plots representing the full range of yellow-red tropical soils and the MDFs that dominate the lowland Bornean landscape. In addition to recording representative forest profiles along transects, at sites of uniform geology, I had suspected that much of the floristic variation was influenced by soil fertility, perhaps individual nutrient ions, as in temperate forest communities. Soil analytical laboratories hadn't been available in Brunei, but in Sarawak, facilities were available. So, this time around, we sampled soils: at the surface, where organic duff was concentrated, and at a standard depth of 30 inches (75 centimeters), where tropical soils would be mineral alone, bereft of visible humus. But bringing back these heavy samples was a nightmare, not least because it had to be done without delay, before microbial activity influenced decay rates and the release of nitrogen and other salts into Otherwise, in those initial years, camp life during surveys became routine, with little of excitement to report. One exception was the accident our team experienced on the upper basalt slopes of Bukit Mersing, a mountain in central Sarawak where thirty plots were eventually installed, four of which became permanent for periodic recensusing. The Bornean climate is almost windless, except 00for the squalls that foreshadow the frequent afternoon thunderstorms. Occasionally, these take the form of violent cold-air downdrafts, flattening the forest in a patch of fifty acres or more. Our camp found itself in one when a giant emergent tree toppled nearby. The team hid beneath its cylindrical trunk, while the camp itself, including the tent frame and its tarpaulin, were trashed. But nothing was lost, and the work could continue. And there were the occasional culinary surprises. We discovered, to our amazement, that civets at some camps would bite into the cans of tinned mackerel and suck out their contents. How did they know what was within? We deduced that a smell, or dry juice, had been left outside during the canning process. And, then again, there was the jaoung: Bornean forests have few canopy palm species, and most While the mixed dipterocarp forests of Borneo had long been viewed as a random assortment of species, discernible and predictable patterns emerged through Ashton's research. Here, a tree climber collects fruits from Sterculia megistophylla (mallow family, Malvaceae) within a research plot at Ulu Mujong. are scattered or local. This one is a Pholidocarpus, which locally formed small groves in damp valleys. It would be felled on discovery and cut open to reveal the massive starchy pith with the portly grubs of a large beetle nestled inside. The trick was to pick these up by the head and bite off the wriggling body, which resembled a greasy polythene tube full of shortening. Then, you had to swallow fast while the three pairs of scratchy legs tickled past your uvula. For me, once was enough! We ultimately conducted four recensuses on five-year intervals, but the work still continues as I write. Different rainforest species achieve trunk-diameter growth rates from ten millimeters to less than two millimeters per annum, which is similar to a stand of regenerating red oaks (Quercus rubra) in a Massachusetts forest. The majority of individuals in any mature MDF, however, are losing to competitors and in slow decline. It required twenty years to gain sufficient data to start comparing the dynamic performance of forests on contrasting rock and soils, and then infer potential lengths of felling cycles and means of sustainable management for timber. By that time, I had spent twelve years on the biology faculty of the University of Aberdeen, where tropical forest research of another kind was occurring, and the work continued after I joined the Arnold Arboretum in 1978. It would take longer still before Harvard graduate student Matthew Potts was to use our data in his doctoral dissertation and Peter Ashton in Northwest Borneo 37 primarily at eight sites, occurred on sandy soils, characterized by higher acidity, lower nutrient levels, and a distinct surface layer of slowdecomposing raw organic matter. The other forty-eight plots, mainly at the other five sites, occupied the more widespread fertile loams, with a higher capacity to retain water thanks to their open lattice of clay molecules. These major groups were consistent, regardless of the underlying bedrock (sedimentary or igneous). Based on Matthew's dendrogram, it could be argued that the reason why two-thirds of the species differed in our original Bruneian plots was simply because the distance between the two sites--sixty miles--is sufficient for the random turnover that might be expected over time from their restricted seed dispersal distances. But the plot sites from Sarawak correlated with soil properties, irrespective of their location, indicating that the selective influence of the physical habitat dominates the random effect of seed dispersal over time in structuring the tree communities. This is particularly evident for isolated plots, where the soil type differs from the surrounding forest, supporting a floristic island of tree species that differ from the surrounding habitat sea. Immediately, I real- NYABAU North Central 0.4 IJU HILL North Central Dissimilarity Index 0.5 0.6 NORTHERN LAMBIR HILLS Northeastern SANTUBONG Southwestern SEGAN North Central BOK-TISAM Northeastern SOUTHERN LAMBIR HILLS Northeastern RAYA HILL South Central MOUNT MERSING North Central BAKO Southwestern ULU BAKONG Northeastern 0.7 NORTHERN LAMBIR HILLS Northeastern CARAPA PILA South Central 0.8 0.9 ULU MUJONG South Central BOK-TISAM Northeastern ULU DAPOI Northeastern ULU DAPOI Northeastern SOUTHERN LAMBIR HILLS Northeastern Taxonomists create phylogenetic trees to show the relatedness between organisms. Ashton, Potts, and colleagues used a similar visualization to classify 105 forest plots in Sarawak. This dendrogram groups the plots according to nine physical habitat characteristics (including altitude, steepness, and soil chemistry). The major branch on the left includes forty-eight plots on fertile loams; the branch on the right includes fifty-seven plots on sandy soils. ADAPTED FROM POTTS ET AL., 2002 devise the most informative methods to analyze patterns and correlations. But even the first results were impressive. Our 105 plots covered 157 acres and included just under two hundred dipterocarp species exceeding one-foot girth. This amounted to four-fifths of all dipterocarps known from Sarawak and Brunei, even though our plots only included MDFs and therefore excluded the characteristic species of the peat swamp forest and other specialized communities. We even captured 70 percent of approximately thirtyfive hundred known tree species in all families. This is tribute to our careful selection of sites representative of the full range of predicted species-specific habitats. These initial results were amplified by Matthew, who devised an elegant method of visually depicting the similarities (or dissimilarities) between the plots. The model, known as a dendrogram, resembles a family tree, with the twigs and branches grouping plots according to their similarities. His analysis revealed several patterns of importance. First, the plots were grouped into two major branches, confirming the original observations and ordinations from Brunei: Fifty-seven Sarawak plots, 38 Arnoldia 77\/1 ? August 2019 Ashton's research had important implications for sustainable forestry, as well as conservation. Similajau National Park, in northeastern Sarawak, received national protection in 1976. ized that the limited dispersal of pollen and seeds results in tropical tree species evolving more-restricted, habitat-defined spaces through competitive speciation. These floristic islands form the ecological equivalent of terrestrial archipelagoes. Other organisms such as symbiotic fungi or insects whose larvae specifically depend on those trees will be similarly confined. The implications for conservation planning are obvious. But although the plots within a particular site occupied distinct terminal branches of the dendrogram (no plots were identical), plots from a particular site generally remained within a single subsection of the dendrogram. This was even true for neighboring sites like the Bok-Tisam Forest Reserve and Ulu Bakong, which are located about twenty miles apart in northeastern Sarawak. Even though these sites share geology and soil characteristics (both primarily occur on yellow-brown loam), their forest compositions could be distinguished from one another. This provided strong evidence that, whereas soil properties dominate at broad landscape scales, the effects of dispersal are dominant over shorter distances, provided the Peter Ashton in Northwest Borneo 39 soils are uniform. Within a single site, we also found that the floristic composition of the plots correlated with topography, as well as nutrient concentrations in the soil. Topography and nutrient concentrations were themselves interrelated, but we were eventually able, by clever analysis, to find that nutrient influences are generally stronger. This research ultimately provided the basis for sustainable management of Bornean MDFs for timber production. Foresters in Peninsular Malaysia have never conducted plot surveys aimed at defining floristically distinct forest types, but they had, over many decades, developed a means of sustainably harvesting timber from MDFs by simulating natural succession: The forest is clearcut in patches hardly larger than large windthrows, after first checking to confirm that there was sufficient natural regeneration that would survive the logging process. Felling cycles of fifty to seventy years were envisaged. Our recensuses confirmed that Bornean forests on loam soils could be managed using the same method. But foresters from Peninsular Malaysia are still finding it difficult to develop sustainable management systems for their high-hill and coastal dipterocarp forests, which floristically resemble the Bornean MDFs of low-nutrient sandy soils. Our findings have suggested a more sophisticated management system is necessary for these forests. Our forest structure measurements and recorded growth rates on these indicated that species attaining timber diameters were slow growing and in lower numbers, although there was an adequate cohort of young trees, rather than seedlings, for successive crops. Sustainable management would, therefore, require foresters return to the same stand at shorter intervals to selectively harvest individual trees, given that it might take a century for a seedling to mature to timber size. For this method, a more experienced workforce would be essential. Our research also supported a new protocol for identifying the locations--to be tested by field censuses--of areas with potentially outstanding species diversity or concentration of endemics, meriting strict conservation. In this way, previously unexplored areas of conservation importance can be identified due to their surface geology. Up to then, conservationists were unable to make such extrapolations. Three national parks in Sarawak had already been legislated in the 1960s. I proposed five more, aimed at comprehensive representation of the flora. These were successfully passed into legislation by the new government of an independent Sarawakian state within Malaysia, in the 1970s, long after our departure from those raucous longhouse days and our entrance into the aethereal realm of academia. Additional reading: Ashton, P.S. 1964. Ecological studies in the mixed dipterocarp forests of Brunei State (Oxford forestry memoirs, no. 25). Oxford: Clarendon Press. Ashton, P.S. 1982. Dipterocarpaceae. In C.G.G.J. van Steenis (Ed.), Flora Malesiana (Series 1, vol. 9, pp. 237?552). The Hague: Matinus Nijhoff. Ashton, P.S. and Hall, P. 1992. Comparisons of structure among mixed dipterocarp forests of northwestern Borneo. Journal of Ecology, 80: 459?491. Ashton, P.S. 2014. On the forests of tropical Asia: Lest the memory fade. Richmond, Surrey: The Royal Botanic Gardens, Kew, in association with the Arnold Arboretum of Harvard University. Beccari, O. 1904. Wanderings in the great forests of Borneo. F.H.H. Guillemard (Ed.) and E.H. Giglioli (Trans.). London: Archibald Constable and Co. (Original work published 1902.) MacArthur, R.H. and Wilson, E.O. 1967. The theory of island biogeography (Monographs in population biology). Princeton: Princeton University Press. Potts, M.D., Ashton, P.S., Kaufman, L.S. and Plotkin, J.B. 2002. Habitat patterns in tropical rain forests: A comparison of 105 plots in northwest Borneo. Ecology, 83(10): 2782?2797. Peter Ashton is Harvard University Bullard Professor Emeritus and was director of the Arnold Arboretum from 1978 to 1987. Among many career honors, his research on tropical forests was recognized with the prestigious Japan Prize in 2007. He and his wife, Mary, live in Chiswick, London. The map in this article was created using Esri, HERE, Garmin, (c) OpenStreetMap contributors, Sources: Esri, USGS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, Intermap and the GIS user community. "},{"has_event_date":0,"type":"arnoldia","title":"Leaning into Legend: Acer mono","article_sequence":4,"start_page":40,"end_page":41,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25666","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070ab26.jpg","volume":77,"issue_number":1,"year":2019,"series":null,"season":null,"authors":"Hetman, Jon","article_content":"HETMAN, J. 2019. LEANING INTO LEGEND: ACER MONO. ARNOLDIA, 77(1): 40 Leaning into Legend: Acer mono Jon Hetman A mbling over nearly every acre of the Arnold Arboretum over the past two decades has made many of its pathways and collections special to me, but perhaps my favorite journey of all is the one I take nearly every day from the Forest Hills Gate to the Hunnewell Building--the lion's share of my morning commute. My great fortune in walking the short mile from home to work in a city recently named the nation's worst for rushhour traffic* is certainly not lost on me--nor is the fact that it is always a journey of beauty and discovery as the seasons ebb and flow. One tree I never tire of admiring along my way is a superb accession of painted maple (Acer mono, accession 5358*A) leaning somewhat languorously toward the road and welcoming visitors to our national collection of maples. Walking in either direction down Meadow Road, the tree is nearly impossible to miss and even harder to ignore. Though not extremely tall--Acer mono typically tops out at a moderate thirty to forty feet at maturity--it has a beautifully low, broad, and symmetrical crown that suggests the kind of idealized form that bonsai artists pursue. In spring, yellow-green flowers appear in tandem with the pale green foliage, which in autumn may turn a yelloworange or apricot. Come winter, the tree's graceful architecture shines through its bare canopy, and the sight of sparkling snow twisting across its branches never fails to send me running for my camera. And then there's the soft twist and southward tilt of the trunk, subtended by a thick knot of exposed roots seemingly coiled like a snake at its base. These thickly layered roots spread out in the opposite direction of the lean, illustrating how tree structure leverages the dynamics of tension and compression (like a suspension bridge) to mitigate the gravitational forces that might otherwise topple them over. While the case can be made that this individual represents its species rather well, the taxonomy of Acer mono remains largely unresolved. Although the Arboretum recognizes A. mono as the correct name, authorities have been mixed on both the name and identity of this widespread maple, often placing it within A. pictum, among other taxa. Seed for 5358*A was received in 1902 from the Imperial Botanic Garden in Tokyo, an institution that has shared material with the Arnold Arboretum since Charles Sprague Sargent made his pioneering expedition to Japan in 1892. Painted maple inhabits the forests of Japan, and it can also be found in China, Korea, Mongolia, and eastern Russia. The Arboretum's wild-collected holdings of the taxon include material collected on several North America-China Plant Exploration Consortium expeditions, including the 2018 expedition to western Hubei Province. Nevertheless, this particular tree has attaind a level of celebrity at the Arboretum. Its position at the head of the maple collection, contrasting handsomely with the texture and seasonal hues of the showy Japanese (Acer palmatum) and Korean (A. pseudosieboldianum) maples nearby, contributes to its appeal. As such, the tree and its neighbors receive attention in many of the Arboretum's public tours each growing season. When famed horticulturist Michael Dirr published the 1983 edition of his seminal Manual of Woody Landscape Plants, following his tenure as a Mercer Fellow at the Arboretum in 1979, he heralded this painted maple as \"one of the most beautiful trees in the Arnold Arboretum.\" And this praise has unwaveringly remained in subsequent editions. This individual may, in fact, be the most famous and recognizable painted maple in the world: as of this writing, a photograph of it adorns the entry for the species on Wikipedia. Its status as a botanical treasure and museum object, one perhaps endangered by an overabundance of public attention, has been acknowledged more definitively of late by the Arboretum as well. Like other eminent accessions across the Arboretum, the tree has been roped off to preserve the health and integrity of its root system from soil compaction. Hopefully this additional protection will contribute to its well-being for years to come and allow this noble maple to be admired as a masterpiece for as long as nature wills. Jon Hetman is the associate director of external relations and communications at the Arnold Arboretum. *INRIX 2018 GLOBAL TRAFFIC SCORECARD "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23466","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14e8128.jpg","title":"2019-77-1","volume":77,"issue_number":1,"year":2019,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Searching for Wilson's Expedition to Australia","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25662","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060896b.jpg","volume":76,"issue_number":4,"year":2019,"series":null,"season":null,"authors":"Grose, Margaret","article_content":"GROSE, M. 2019. SEARCHING FOR WILSON'S EXPEDITION TO AUSTRALIA. ARNOLDIA, 76(4): 2?13 Searching for Wilson's Expedition to Australia Margaret Grose C ould diaries, newspaper clippings, and letters be hidden at the Arnold Arboretum, unexamined for almost one hundred years? Might the Arboretum possess more than two hundred glass plate negatives by famed plant collector Ernest Henry Wilson without labels for location or species? In the spring of 2016, I met Arboretum scientist Peter Del Tredici while visiting MIT, and he invited me to see the Arboretum, an invitation I accepted with relish. After a tour of the grounds, including a walk in the woods, a stop on Peters Hill, a discussion of insect attack on hemlocks, and a look at the amazing old bonsai, we found ourselves in the horticultural library. \"Bet you Australians don't know that Wilson went to Australia in the 1920s,\" Peter said. \"No one has looked at the collection. It's sitting there.\" He pointed, and there it was, hidden in plain sight. When I returned to Australia, I inquired as to whether, indeed, botanists knew that Wilson, famed for botanical explorations in China, had travelled to Australia. No one did. Everyone that I spoke with was astounded. Herbaria staff did not know; botanists at my university and elsewhere did not know; even those who have a long and keen interest in the botanical exploration of the Australian continent did not know. Wilson's expedition between 1920 and 1922 was to Commonwealth countries and no others. During my time in the United States, I have been asked, on a number of occasions, what the Commonwealth is, what countries belong to it, and what it means. The Commonwealth consists of now-independent countries that once made up the British Empire, and these countries are joined by commonalities, such as a politically independent judiciary and public service, school uniforms, and cricket, the second-most popular sport in the world after soccer. Wilson's travels took him to countries now known as Australia, New Zealand, Malaysia, Singapore, India, Sri Lanka, Kenya, Uganda, Zimbabwe, Zambia, and South Africa, before returning to England and then on to Boston. He wrote about the horticultural aspects of this trip in various garden magazines, as well as in the first volume of his book Plant Hunting, published in 1927 (and reprinted decades later as Smoke that Thunders). The writing in the book is part tourist travelogue and part horticultural journal, with a discussion of the cultural requirements and garden potential of plants met along the route. What surprised me was how much Wilson knew about the discovery of the western coast of Australia by the Dutch in the early 1600s, sailing eastwards from Cape Town to use a faster route to the East Indies. This is a lesser-known history than that of Captain James Cook mapping the east coast of Australia in the 1770s, and it shows that Wilson was remarkably well-informed in 1920. Two years after learning about Wilson's expedition, I returned to the Arboretum with a Sargent Award for Visiting Scholars, intent on examining the archives, images, and herbarium specimens pertaining to Wilson's time in Australia. I was fascinated by the idea of a lost collection of Australia and, like many botanists, interested in the history of plant collecting. I was also curious as to whether the images might offer a glimpse of land now lost to agriculture and suburban development, both research areas of my own. What I found in the archives was a collection in excellent order, with annotated boxes, carefully arranged glass plate negatives that had been digitized in 2018, meticulously kept diaries with their labels and individual folders in boxes, and neatly ordered newspaper clippings, letters, and ephemera of the expedition. I took photos of every box and every label. It was not evident who had established the organization of the collection, but even with the clear organization, there was little documentation to suggest what was in the letters, or what the images showed and where they were taken. PHOTOS FROM ARNOLD ARBORETUM ARCHIVES UNLESS NOTED E.H. Wilson in Australia 3 In Plant Hunting, Wilson described eucalyptus like these massive karri (Eucalyptus diversicolor) as \"Australia's noblest trees.\" Karri are considered the second-tallest flowering tree species in the world, after E. regnans of southeastern Australian. Note the numerical label: Y-347. 4 Arnoldia 76\/4 ? May 2019 Charles Lane-Poole--formerly unidentified in Wilson's collection of glass plate negatives--stands beside a massive eucalyptus. In a letter to the Arboretum, Wilson mentioned that, soon after arriving in Australia, Lane-Poole \"became guide, philosopher & friend.\" E.H. Wilson in Australia 5 While images from India and Africa were annotated with old typewriter notes on thin white paper, the Australian images--numerically numbered and coded with the letter Y--were not annotated, and it was unclear just where Australia started and ended. It became clear to me that my inquiry would require three components: The examination and transcription of diaries, letters, and notes; the annotation of the unlabelled images; and a search of the Arnold Arboretum Herbarium to see what plant specimens had been submitted there by Wilson's expedition. Wilson arrived in the port of Fremantle, on Australia's western coast, after a tedious and hot trip by boat from Sri Lanka (then Ceylon). He was immediately struck by how Perth, the capital of Western Australia, reminded him of southern California. (There is a saying in Perth, my hometown, that we have the climate that the Californians think they have.) Southwestern Australia has, like southern California, hot summers and mild winters, and this was important to Wilson because Henry Huntington, a sponsor of the expedition, was eager to hear of plants suitable for cultivation in California. This region, today referred to botanically as the Southwest Australian Floristic Region, is known for its exceptionally high levels of floral endemism--meaning plant species that are found nowhere else in the world. Wilson was astounded by the plants he found there. \"To a visitor from the Northern Hemisphere, no matter how familiar he or she may be with the forest scenery of the North, Western Australia is a new world,\" he wrote in Plant Hunting. \"Nay, it might well be part of another planet so utterly different is the whole aspect of its vegetation. Intimate knowledge of the plants of the boreal regions only serves to accentuate the variance.\" i While reading Wilson's thoughts on the Australian flora, I was struck by his observational clarity, especially after recently reading a paper on the evolutionary history of the Australian flora over the last sixty-five million years. \"To what extent is the Australian flora unique?\" the authors--botanists Michael Crisp and Lyn Cook--asked, perhaps asking because most Australians think it is and most foreigners agree with them. They noted that thickened foliage (sclerophylly) is not unique to Australia, nor are traits that promote water storage (xeromorphy), nor is fire as a selective genetic force unique to the continent. But nowhere else do these combined traits dominate large proportions of an entire continent. ii Wilson, however, immediately noted two traits as completely unique to Australia: the angle of leaf repose and the color of the flora. \"In the North our trees in general have spreading umbrageous crowns, dark, often lustrous green leaves which ... cast a heavy shadow,\" Wilson wrote. \"In Western Australia the dominant trees have open, tufted crowns, gray or glaucous green leaves which ... cast little or no shadow. This difference in the color of the treefoliage and the fact that the leaves are pendant instead of spreading on the branches may seem to the reader trivial matters, but in reality they completely change the aspect of the forests and profoundly influence the whole landscape.\" iii To have quickly put his finger on these two points is a tremendous perception. The color of the Australian flora presents a world of very different greens and grays to the Northern Hemisphere, as I have found in my own research, and Wilson appeared to have recorded them all. The unique flora of Australia made it relatively easy for me to tease out Australian images from the others. I also began to identify the unnamed individuals in the photographs. One man, who recurred in multiple photographs, was distinctive due to a hook in place of his left hand. I emailed Australian colleagues about a botanist matching this description (which I thought would have been easy), but I met a blank. Eventually, I came across a biographical entry about Charles Lane-Poole, the conservator of forests in Western Australia from 1916 to 1922, which mentioned that his left hand had been lost in a shooting accident when he was nineteen (all other biographical sketches politely did not mention this). I had my man. I later came across Wilson's comment that he had travelled two thousand miles with LanePoole \"through all the important forest areas.\" Elsewhere, in \"the sand plains and savannah regions,\" he was guided by Desmond Herbert, 6 Arnoldia 76\/4 ? May 2019 the Western Australian government botanist, who was also shown in images. Wilson noted that, without such expert guides, \"I should have been completely lost among the extraordinary varied and anomalous vegetation.\" These men were top in their field and their companionship and assistance shows the esteem in which Wilson and the Arnold Arboretum were held. The images, however, revealed an oddity--all that I could readily identify were from the beginning of his trip in Western Australia. With limited time, I set to work, realizing that diaries could be transcribed, letters read, and images annotated in Australia. Thus, I headed to the Harvard University Herbaria, where the Arboretum Arboretum's wildcollected herbarium specimens are housed, because I recognized that those materials could not be examined later. While Wilson's notebooks and photographs would provide some sense of his route across the continent, the specimens in the herbarium would provide more detailed information about where and when Wilson travelled, what he was collecting, and the identities of other botanists who collected on behalf of the Arboretum. At the Harvard University Herbaria on Divinity Avenue, I was given pencils, little paper envelopes for broken off bits of specimens, and clear plastic clips to replace rusty metal clips from one hundred years ago. The collection is contained within rolling cabinets, called compactors, and I was presented with a written directive that \"compactors can be opened by releasing the locking bar and smoothly rotating the handle of the appropriate bay.\" The instructions also suggested to \"move one row at a time to prevent strain on the system,\" although I was not sure whether that was referring to the compactors or to me. Lists of plant families hung on the walls, indicating where the families were housed in the building's several floors and annex rooms. \"Where would you like to start?\" Anthony Brach of the herbaria's curatorial team asked me. I looked about wideeyed. Well, why not Myrtaceae? I ventured, and Anthony chuckled. I had chosen \"the big one\"--the family of the eucalypts. Eucalyptus is a major genus in Australia and is rare in the world in that this single genus virtually defines an entire continent. There are now over eight hundred species (the number always rising), and this usually surprises overseas visitors to Australia, who often think that there are only a few \"gum trees.\" The eucalypts include the world's tallest flowering plant, E. regnans, known as the mountain ash, which reaches heights of 295 feet (90 meters). Timber records in the nineteenth century reported that trees logged then had reached far greater heights--up to 490 feet (150 meters). This loss underscores one of the reasons Wilson was keen to see the Australian forests. Eucalypts, however, are not all tall; some are small and gnarled; many are low shrubs and suitable for home gardens; some have brilliant flowers; some have dusky gray leaves that are suitable for the flower trade; some possess huge bud caps that gave the genus its name-- eu (well) and kalyptos (covered); some are small and gnarled; some are single-trunked; others have a mallee form, an Aboriginal term referring to plants with multiple trunks that emerge from underground lignotubers. Most live in mixed woodlands. Wilson was captivated by their variety and the colors and forms of trunks. He noted, for example, that the salmon gum (Eucalyptus salmonophloia) \"is a handsome tree, with a smooth white to pinkish trunk ... the twigs are reddish\" and that the gimlet (E. salubris) is \"fluted and twisted like a screw-- hence the name gimlet.\" iv In photographs, he recorded fire-scarred trees, and he was clearly impressed by the extraordinary capacity of most eucalypt species to resprout after fire. During the next few weeks, I extracted hundreds of specimens from the expedition across a large range of families and genera, all from blue folders that indicated the specimens came from Australia. These were easy to spot among swathes of green, orange, pale orange, yellow, pale yellow, beige, white, off white, and ranges of pinks and reds. In short, if a folder was blue, I knew it was mine to take out and examine. How curious and exciting it was to search through the blue folders and find labels with Wilson's name and handwriting. \"Near a saltlake,\" he noted on one. \"In a group of trees,\" he wrote on others. He described a now-rare little shrub (Daviesia euphorbioides) as a \"centipede bush\" and called a large shrub (Banksia sessilis) SPECIMENS FROM HARVARD UNIVERSITY HERBARIA Herbarium specimens provided historical evidence about Wilson's travels through southwestern Australia. Clockwise from top left: Hakea platysperma, a small shrub, collected on the sandplain at Yoting on October 25, 1920. Casuarina fraseriana (now Allocasuarina fraseriana) collected at Albany, on the southern coast, on November 6. Eucalyptus ficifolia (now Corymbia ficifolia) collected from the Frankland River region in November (day unspecified). Eucalyptus flocktoniae collected near Widgiemooltha in November (day unspecified). SPECIMENS FROM HARVARD UNIVERSITY HERBARIA 8 Arnoldia 76\/4 ? May 2019 Diverse members of Proteaceae include: Banksia grandis (left) collected October 25, 1920, in the Darling Scarp, east of Perth. Wilson noted the yellow flowers--produced in a large cone-shaped structures--were collected from a small tree, about 15 feet (4.5 meters) tall. Hakea multilineata (right) collected 20 miles (32 kilometres) south of the oncethriving gold-mining town of Coolgardie, now a near ghost town. \"parrot bush.\" Many labels included collection numbers that related to the diaries. From the images, I knew that Wilson and his travel companions appeared to have gone by horse, train, and a roofless Model T Ford. They travelled south of Perth through wetlands near Busselton, into the Darling Scarp--the hills east of Perth--to collect in the jarrah (Eucalyptus marginata) forest. They continued south to the tall karri (E. diversicolor) forests, east through heavily cleared land of wheat and sheep fields, and still farther east into semi-arid country. Labels in the herbarium revealed place names: Yoting, Burracoppin, Merredin, Toodyay, Widgiemooltha, Quairading, Coolgardie, and Kalgoorlie. Many of these are indigenous names given to country towns or settlements that in 1920 and 1921 were gold-mining camps. At one such now-tiny outpost, Westonia, Wilson noted in his diary on October 22, 1920, that \"Saturday night in a mining camp is not a quiet place. Singing and chat extended far into the night.\" Looking at the labels in the herbarium, I was amazed to see where Wilson went--to places remote enough now, let alone then. A whole suite of specimens was collected \"20 miles south of Coolgardie,\" in hot and dusty conditions, but Wilson enthused over the \"astonishing\" variety of species. In his diary, he wrote, \"they are mostly prickly in character and many of them especially so.\" v Reviewing the specimens belonging to the Proteaceae was a treat for me because my doctoral dissertation had been on the ecophysiology of Banksia, which is the most well-loved member of the family in Australia. As I opened the compactors, I could see the banksias in their blue folders, and I laughed--they were large, MAP BY ARNOLD ARBORETUM AND GIS COMMUNITY E.H. Wilson in Australia 9 Western Australia Kalgoorlie Coolgardie Burracoppin Merredin INDIAN OCEAN Toodyay Cunderdin Northam Mundaring t York W h e a PE RTH Fo re Jarrahdale nnah Bruce Rock Darwin Northern Territory K rr a Donnelly River Queensland Western Australia iF o re st Pemberton Brisbane South Australia PERTH s Walpole New South Wales Frankland River Denmark Nornalup 50 MILES Lake Lefroy Widgiemooltha Capel Busselton ah Ja Cookernup Totadgin belt Sava Pingelly rrah North Dandalup Moorine Rock Yoting Beverley Quairading sts Fremantle S a vann Westonia Hines Hill Sydney Albany Frenchman Bay Adelaide Victoria Canberra Melbourne Tasmania SOUTHERN OCEAN Hobart 100 KILOMETERS Locations recorded on Wilson's herbarium specimens and in notebooks revealed extensive collections throughout southwestern Australia, from the towering karri (Eucalyptus diversicolor) forests, south of Perth, to the semi-desert forests farther east. Contemporary train lines are shown in gray. The Trans-Australian Railway, completed in 1917, connected Kalgoorlie with eastern cities. chunky folders. Banksia flowers grow in hairy cone-shaped inflorescences, which measure up to seventeen inches long. Some of the cones-- the flower heads or remaining fruiting bodies-- were stored elsewhere, in boxes, due to their size, and Anthony Brach got them out for me. These attracted comment from passing taxonomists in the herbarium, but I pitied that they could not see them on the trees instead of these dead, dry relics. One even commented: \"How wonderful to work on these. Some people,\" she shuddered, \"have to work on grasses!\" Wilson collected various banksia species near Perth and inland, and many of his images are of banksia trees. As I pulled out specimens, I mused as to what Wilson must have thought about when he looked at the banksias, with their wire-tough thick leaves, erect demeanour, and vibrantly colored inflorescences, usually buzzing with insects and birds with tongues and beaks evolved to suck nectar from these exact trees. In Plant Hunting, he described the genus as \"among the most wonderful flowering trees of Australia.\" He described the large pale lemon and yellow flowers of Banksia grandis, with cones measuring up to eight inches long, and he noted elsewhere that the species should be grown in California. Beyond the visual appeal, many contemporary botanists were intrigued by the banksias--which are particularly prominent in southwestern Australia--because these were clearly related to the striking diversity of Protea from South Africa. Our contemporary understanding of continental drift was first proposed by the German scientist Alfred Wegener in 1912, and we now know that Antarctica, Australia, South America, Africa, India, and 10 Arnoldia 76\/4 ? May 2019 Kingia australis--slow-growing and long-lived monocots--fascinated Wilson. He called them \"weird and extraordinary\" and suggested, in Plant Hunting, that \"the scene would be more complete if ... a Nothotherium, a Dinosaurus, a Plesiosaurus or some other monster of the remote Lizard Age\" were present. E.H. Wilson in Australia 11 Wilson returned to the Arnold Arboretum in 1922, after visiting India and various Commonwealth countries in southern Africa, which were recorded in his images. But what had happened to the herbarium specimens and images from a major part of his Australian tour? One day, in the middle of my stay, I turned to materials in the archives, to see if I could shed some light on the mystery of the missing Australians. The key was in a box of newspaper reports. An article described Wilson's \"great disappointment\" at losing much of the collection because the boat carrying two large consignments of photographic plates and specimens was lost at sea. The ever-helpful Lane-Poole had earlier shipped the entire Western Australian collection, which arrived safely in Boston. Sargent's response is not known. He had considered a study of Tasmanian conifers \"one of the most important objects of the expedition,\" vi but those collections were all gone. Other questions were answered: printed black-and-white images from Tasmania were sent by the noted Hobart photographer John Beattie after the loss of Wilson's collection, and unlabelled lantern slides, sent to Wilson, all depict New Zealand in a travelogue style. New Zealand were once joined in the supercontinent Gondwana and shared flora. Proteaceae is one of the oldest families in the Southern Hemisphere and once was found across Gondwana, right across what is now Antarctica. Fifty-million-year-old fossils have been found in Patagonia and New Zealand that even an untrained eye would easily recognize as lovely banksias. But alas, as Wilson knew, none of these living beauties would grow in Boston's climate for the Arboretum. As my notes from the herbarium accumulated, I soon realized that, as with the photographs, the specimens were almost entirely collected in Western Australia. This was odd, as Wilson's diaries revealed that he had travelled by train across Australia after leaving Perth, and had visited Adelaide, Melbourne, and Sydney, where he spent Christmas in 1920. He then went into the New South Wales heartland, north into Queensland, and then eventually to Tasmania in April 1921, after a sojourn in New Zealand. The diaries said little but did give the names of plants collected, with collection numbers. Yet, while I came across the odd specimen from Queensland and Tasmania, I found none collected by Wilson from New South Wales or Victoria. This was a mystery. At the end of each day in the herbarium, which closes precisely at five o'clock, I would close the compactors, pull down the bar, and turn off the light, as directed. One of the many delights of my own expedition into the 1920s expedition was the people I met along the way who still enliven the images and specimen sheets. Many of the herbarium specimens had been reassigned by David Moresby Moore, a British plant taxonomist and systematic botanist with a major interest in the flora of Gondwana, Patagonia, and the Antarctic and sub-Antarctic islands. His work amid the Australian collection at Harvard in the 1960s made my job far easier, because the plants were correctly assigned and thus correctly catalogued. \"DM Moore\"--his appellation on the bright white labels near Wilson's originals-- was a welcome sight. Other collectors assisted the Arnold Arboretum and donated specimens from Western Australia, South Australia, and Queensland as part of the Expedition. Most important were those submitted by Frederick Schock (\"F.M.C Schock\" on the labels), who was a forest ranger in Western Australia. Schock's specimens were largely collected in 1916, but they were sent to Harvard after Wilson returned and were labelled \"Arnold Arboretum Expedition to Australasia, India, and Africa.\" Many included flower parts that had not been seen by Wilson during his summer visit, because many of these species, typical of southern Australia, do not flower in summer. With the addition of these donated specimens, one of the missions of the expedition began to become apparent to me: to add to the global collection of specimens and images held at the Arnold Arboretum. The Arnold Arboretum celebrated its fiftieth anniversary in 1922, and Charles Sprague Sargent, the founding director, laid out this global vision in a report on the Arboretum. \"If the Arboretum is to become 12 Arnoldia 76\/4 ? May 2019 Wilson stands within a jarrah (Eucalyptus marginata) forest, in the Darling Scarp, east of Perth. The trees beside him have been scarred by fire. a great institution for gathering and spreading information about trees and allied plants, specimens and a series of photographs of every species of tree in the world should be found in the herbarium,\" Sargent wrote. His ambition was that the work already achieved \"should be extended over the rest of the world.\" vii Given Sargent's desire to develop global collections of photographs and herbarium specimens, it made sense that additional Australian specimens would be requested from Schock and others, to fill gaps within Wilson's collection. While the Arnold Arboretum could only grow plants that suited Boston's cold winters and humid summers, Wilson's expedition to Commonwealth countries revealed the equal importance of the herbarium and archives for institutional collecting goals. I had noted, in the herbarium, hundreds of specimens from Australia that were not from trees and that were of botanical interest, rather than of forestry value, reflecting Sargent's desire for a comprehensive collection. I was also struck by letters of thanks from Sargent regarding the donation of pamphlets and books on specific aspects of the world's flora, as well as the contribution of physical specimens (including cones and seeds), sent from the countries visited on Wilson's expedition, even though these would not grow in Boston. With these materials, the various missions of the expedition to Australia became even more apparent. In addition to collecting images and specimens, Wilson was to make connections between the Arboretum and the staff of other international botanical institu- E.H. Wilson in Australia 13 tions, while investigating potential timber trees for production in the United States and assessing firsthand the state of forests in the world. Wilson achieved all these goals despite the loss of much of the physical collection. Wilson's travelling companions in Western Australia were highly informed and are now famous men in Australian botany. I wondered what they talked about, especially given Lane-Poole's desperate unhappiness with the lack of forest protection in Australia, and Wilson's diary comments about the ruthless destruction of woodlands for agriculture in Australia.viii Only a few months after Wilson's visit, Lane-Poole resigned, in 1922, as the conservator of forests because the government did not appear interested in conservation but was solely concerned with timber extraction. Letters between Wilson and Sargent show that the whole idea of forest protection was of great importance to the expedition because they both saw that forests were under threat across the world. This suggested to me that the conservation movement was more alive in the 1920s than many of us fully appreciate today. Wilson, Sargent, and Lane-Poole all saw that the world's forests were in danger of overexploitation and habitat loss, and both Wilson and Lane-Poole named the loss of large old trees as of greatest concern in Western Australia. Yet one hundred years later, botanists and conservationists are still raising this issue because it needs to be raised--surely something that would have saddened these men. The great banksia woodlands surrounding Perth have been substantially lost due to suburban development, and the woodlands east of the Darling Scarp, where Wilson noted an abundance of \"curious,\" \"wondrous,\" and \"extraordinary\" plants, continued to be clear-cut for agriculture into the 1980s. Today, ecological agriculture and ecosystem repair are imperatives for the future. With my time at Harvard running out, I had to cease work at the herbarium and say my goodbyes at the Hunnewell Building and the Weld Hill laboratories, where I had my office. And I had found that the answers to my search for the Arnold Arboretum's Expedition into Australia's spectacular flora lay not simply in one place. As Sargent noted, the arboretum is a three-part collection, with a living museum, an herbarium, and a library. ix I had needed all of the resources and staff of the arboretum to begin to understand this last great journey that Wilson undertook. Acknowledgements Margaret Grose thanks the Arnold Arboretum for a Sargent Award for Visiting Scholars, Anthony Brach of the Harvard Herbaria, and Lisa Pearson, Jonathan Damery, Peter Del Tredici, and Ned Friedman of the Arnold Arboretum. The map in this article was created using Esri, USGS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, Intermap and the GIS user community. Endnotes i Wilson, E.H. 1985. Smoke that Thunders. London: Waterstone & Co. Limited. (Original work published as Wilson, E.H. 1927. Plant Hunting, Vol. 1. Boston: The Stratford Company.) iiCrisp, M.D. and Cook, L.G. 2013. How was the Australian flora assembled over the last 65 million years? A molecular and phylogenetic perspective. Annual Review of Ecology and Systematics, 44: 303?324. iiiSee Wilson, 1985, above. ivWilson, E.H. 1920. Alternating diary and collection notes, October 22, 1920 (box 14, folder 3?4). Ernest Henry Wilson (1876?1930) papers, Arnold Arboretum Horticultural Library, Harvard University. v Ibid. viSargent, C.S. 1921. Sargent to E.H. Wilson, April 18, 1921 (volume 9, page 599?600). Charles Sprague Sargent (1841?1927) papers, Arnold Arboretum Horticultural Library, Harvard University. vii Sargent, C.S. 1923. The first fifty years of the Arnold Arboretum. Journal of the Arnold Arboretum, 3(3): 127?171. viii Wilson, E.H. 1920. Alternating diary and collection notes, October 25, 1920 (box 14, folder 3?4). ixSargent, C.S. 1925. The Arnold Arboretum. In Report of the President and the Treasurer of Harvard College, 1923?24 (pp. 229?232). Boston: Harvard University. Margaret Grose is a senior lecturer in landscape architecture at the University of Melbourne, where her research and teaching merges design and ecological science. Margaret has published across science, landscape architecture, and planning. "},{"has_event_date":0,"type":"arnoldia","title":"Deja Vu Viburnums: A World Away but Close to Home","article_sequence":2,"start_page":14,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25660","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060856d.jpg","volume":76,"issue_number":4,"year":2019,"series":null,"season":null,"authors":"Dosmann, Michael S.; Aiello, Anthony S.","article_content":"AIELLO, A.S. AND DOSMANN, M.S. 2019. D?J? VU VIBURNUMS: A WORLD AWAY BUT CLOSE TO HOME. ARNOLDIA, 76(4): 14?25 D?j? Vu Viburnums: A World Away but Close to Home Anthony S. Aiello and Michael S. Dosmann I t is a truth universally acknowledged that closely related plants occur in widely varying locations around the globe, yet going from a truth universally acknowledged to a truth personally observed is a profoundly gratifying process. In September 2018, along Lake Yuno, within Japan's Nikko National Park, we collected fruits of the forked viburnum (Viburnum furcatum). The fleshy drupes were reddish-orange, although a number had already turned blackish and were mushy to the touch. We reached them by standing on tiptoes in the muck, and pole pruners came in handy for those higher up. When not browsed by deer, the shrubs stood up to ten feet tall and sometimes just as wide, the sprawling branches positioning the round, saucer-sized leaves like open hands to capture light in the shady understory of maples and conifers. While this species is native to eastern Asia, we could have fooled ourselves into thinking we were standing in front of the hobblebush (V. lantanoides) of eastern North America, given the striking similarities between the two species' appearance, habitat, and associated genera. It was already a great morning collecting, and not far away, deciduous azaleas (Rhododendron wadanum) and hollies (Ilex geniculata) grew near the shoreline of the mountain lake, as did the occasional clethra (Clethra barbinervis). Higher up on the ridges, large beech (Fagus crenata) and hemlock (Tsuga diversifolia) filled the overstory. Because these genera also occur in eastern North American forests, we could have sworn we had already experienced this collecting day before, a hemisphere away. While we have long understood the profound concept of biogeographical disjunctions, it has taken compounded years of plant exploration for us-- as colleagues at sister institution that share a long history of cooperation and collection goals--to personally observe these connections in far reaching places. Our collaborative expedition to Honshu, Japan, also included Steve Schneider from the Arnold Arboretum, Todd Rounsaville from the Polly Hill Arboretum, and Mineaki Aizawa and Tatsuhiko Shibano from Utsunomiya University. During our two-week trip, we visited six locations and made fifty-eight collections of fifty-five different taxa. Like the viburnum, half of the genera we collected co-occur in eastern Asia and North America, creating even more moments of d?j? vu. Botanical Kin At Lake Yuno, most of the forked viburnum's wide, rounded leaves had heavy insect herbivory, giving it the Japanese common name of mashikari (always eaten by insects), yet the habit and leaf shape instantly reminded us not only of the American hobblebush but another eastern Asian species, Viburnum sympodiale. The genus Viburnum (Adoxaceae) has a wide geographic range, with approximately 165 species occurring across the Northern Hemisphere and dipping into South America and Southeast Asia. Botanists have organized these species into sub-generic groups or clades, based on how closely related these are to one another. These three viburnums occupy the Pseudotinus clade, which also includes a fourth species, V. nervosum (also from eastern Asia). The four share a distinctive branching architecture, which Erika Edwards and colleagues have dubbed the \"furcatum pattern\" (drawing on work Michael Donoghue published in Arnoldia in 1981). The branches grow horizontally with the ground (known as plagiotropism) until the end of the season when one of the terminal buds turns upwards to produce a short, reproductive shoot. The next spring, the main, horizontal growth continues from the other terminal bud, creating a sympodial growth pattern, where one of two forked branches becomes dominant. This branching structure is acknowledged in the specific epithet of V. sympodiale. MICHAEL S. DOSMANN DANNY SCHISSLER MICHAEL S. DOSMANN Viburnum 15 On a 2018 expedition to Japan, the authors and colleagues collected fruits of Viburnum furcatum near Lake Yuno (bottom left). Insects had devoured the leaves (top), yet the resemblance to the North American species V. lantanoides (bottom right) was unmistakable. These disjunct species are two of four representatives of the Pseudotinus viburnums. PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY; PEABODY.YALE.EDU. PHOTO BY PATRICK SWEENEY JONATHAN DAMERY 16 Arnoldia 76\/4 ? May 2019 Unlike the other Pseudotinus viburnums, Viburnum nervosum (left) lacks the ring of sterile flowers around the margin of each inflorescence, although the individual flowers are larger. V. lantanoides (right) shows a more characteristic structure. Although there are no European representatives, this quartet is a fascinating case study of the history of botanical description and horticultural introduction. The description of these four species follows a familiar arc from the initial exploration and cultivation of North American natives in the late eighteenth century, followed by the introduction and study of Chinese and Japanese species through the early twentieth century, and it coincided with an increasingly refined understanding of biogeography and the eventual explanation of lookalike species that are spaced a world away from one another. The native range for Viburnum lantanoides stretches between Canada's Maritime Islands and northern Georgia. This species--the first of the Pseudotinus to be attributed a scientific name--was described as V. alnifolium by Humphrey Marshall, a cousin of botanists John and William Bartram, in an alphabetical catalogue of plants growing in the eastern United States. Marshall's publication in 1785 described the leaves as \"heart-shaped, oval, sharp-pointed, deeply sawed on their edges, strongly veined, and placed opposite upon long slender footstalks,\" but the most telltale characteristic was the halo of sterile flowers encircling each inflorescence--a feature also shared with V. furcatum and V. sympodiale. At first, this portrayal seems unmistakable, but as it turns out, Marshall's description was not of the hobblebush viburnum. Marshall based his description upon an earlier one (by British botanist Philip Miller in 1768), which was confused, in part, with smooth hydrangea (Hydrangea arborescens) given the shared floral characteristics and large leaves that are arranged opposite one another along the stem. Almost twenty years after Marshall's publication, Andr? Michaux proposed the alternative name, Viburnum lantanoides, which is now PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY; PEABODY.YALE.EDU. PHOTO BY PATRICK SWEENEY JONATHAN DAMERY Viburnum 17 While botanists have long considered Viburnum furcatum and V. lantanoides to be closely paired within the four Pseudotinus, recent analyses have shown that Viburnum sympodiale (left) and V. furcatum (right) are more closely allied. widely accepted. (For further background reading on the confusion between the two names, see Ferguson, 1966, and Mackenzie, 1927.) In some sense, it's surprising that such a widespread shrub would have escaped the attention of earlier botanists. After all, one of the shrub's characteristic qualities is its ability to lay branches and root, creating bulky masses of foot-catching vegetation that give the species its common name: hobblebush. When describing the species' habitat, phrases like \"damp woods,\" \"moist woods,\" \"cool moist woods,\" \"rocky woods,\" \"deep woods,\" \"swampy woods,\" and other combinations thereof leap off the labels of most specimens at the Harvard University Herbaria, and when it comes to describing the shrubs themselves, another word dominates: \"thickets.\" Even now, however, the species is scarce in the nursery trade and often slips under the horticultural radar. We first jointly encountered the species near its southernmost range edge in May of 2002, while hiking along the Blue Ridge Parkway in Transylvania County, North Carolina. On either side of the path, the shrubs bore large and roughly heart-shaped leaves that stuck out at wide angles to catch the light. Just two years before, and not far away, Tony had first seen and collected the species in very similar habitats at Nantahala National Forest in North Carolina and near Clingman's Dome in the Great Smoky Mountains National Park. Disjunct Discoveries The next Pseudotinus species to enter the botanical literature was Viburnum nervosum-- the most basal of the four species. While we often think that the earliest Western botanical explorations to Asia occurred in Japan and coastal China, the European presence in India meant that trans-Himalayan species were often described earlier in the nineteenth century, 18 Arnoldia 76\/4 ? May 2019 larger. Showy fertile flowers are an alternative to showy marginal flowers for attracting pollinators, according to Brian Park and colleagues. The Japanese species that we observed at Lake Yuno--Viburnum furcatum--was next to be described. Japan was predominantly closed to Europeans and Americans until 1868, the year of the Meiji Revolution, but in 1858, the German botanist Carl Ludwig Blume obtained an herbarium specimen of a Japanese viburnum and labelled it with this name. The species would not be properly described, by Carl Johann Maximowicz, for another two decades. Nonetheless, in 1859, Harvard botanist Asa Gray noted the presence of a Japanese viburnum that he suspected was the same or a closely related species as V. lantanoides, based on herbarium specimens collected by Charles Wright, botanist on the United States North Pacific Explor- ARNOLD ARBORETUM ARCHIVES PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY; PEABODY.YALE.EDU. PHOTO BY PATRICK SWEENEY before their eastern counterparts. In 1825, forty years after Marshall's putative description of the hobblebush, David Don, the librarian for the Linnaean Society, described V. nervosum, based on collections made by Nathaniel Wallich and Francis Hamilton in Nepal. Neither of us has encountered Viburnum nervosum in the wild, but this species occurs at exceptionally high altitudes and is common in open, boggy habitats at (or even above) the tree line in Sichuan, Xizang, and Yunnan Provinces of China, as well as in Bhutan, India, Myanmar, Nepal, and North Vietnam. The Flora of China notes that it can grow up to 14,700 feet (4,500 meters) above sea level. Unlike the other three Pseudotinus species, V. nervosum does not produce the distinctive halo of sterile flowers around each inflorescence; instead, the fertile flowers within each inflorescence are In 1914, Ernest Henry Wilson photographed Viburnum furcatum growing on a woodland edge in the Yumoto region of Japan, near the location where the authors also observed the species. Among the Pseudotinus, V. nervosum (right) has the most atypical habitat preferences--shown within a montane bamboo and heath scrub at more than 10,000 feet (3,200 meters) elevation in Yunnan, China (notice the broader leaves, tucked in the center of the frame). PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY; PEABODY.YALE.EDU. PHOTO BY PATRICK SWEENEY Viburnum 19 Branching structure is a defining characteristic for the Pseudotinus viburnums. Here, Viburnum sympodiale-- photographed in Hunan, China--shows how the branches fork, with one side continuing the dominant horizontal growth and the other turning upwards to produce flowers and fruit. ing Expedition between 1853 and 1856. This and other early specimens, initially referred to as V. plicatum (and occasionally V. tomentosum), supported Gray's theories of disjunction of species, particularly between the eastern United States and eastern Asia. Gray concludes his momentous essay, published in 1859, with a provocative summation: \"Under the light which these geological considerations throw upon the question, I cannot resist the conclusion, that the extant vegetable kingdom has a long and eventful history, and that the explanation of apparent anomalies in the geographical distribution of species may be found in the various and prolonged climatic or other physical vicissitudes to which they have been subject in earlier times.\" Gray recognized that the occurrence of distant doppelg?ngers like these North American and Japanese viburnums could only be explained through change on a geologic timescale. This argument would gain substantial theoretical weight with the publication, in 1860, of On the Origin of Species, written by Gray's friend and scientific correspondent Charles Darwin. It is noteworthy, therefore, to see this group of Pseudotinus viburnums--or even just the paired Japanese and North American species-- along this timeline of scientific breakthroughs. As botanists like Gray began to work with a greater volume of herbarium specimens from Asia, an evolutionary and geologic story began to emerge. Global exploration and paradigmchanging scientific discovery necessarily moved in tandem. The final viburnum in the quartet--Viburnum sympodiale--was described by Paul Graebner in Ludwig Diels' epic Die Flora von Central-China, published in 1901. Graebner named this species using herbarium specimens MICHAEL S. DOSMANN MICHAEL S. DOSMANN ANTHONY S. AIELLO 20 Arnoldia 76\/4 ? May 2019 Along with ornamental flowers and fruit, the Pseudotinus viburnums produce exceptional fall color. On a 2005 expedition in China, Tony observed Viburnum sympodiale (top) with red fall color. And in the Adirondacks, in 2008, Michael observed a striking mix of colors on V. lantanoides (both bottom). Viburnum 21 collected by Augustin Henry and Arthur von Rosthorn, and he noted how its forked (or sympodial) branching structure resembled that of V. furcatum. Ernest Henry Wilson observed V. sympodiale during his initial trip to China (between 1899 and 1902), while working for Veitch Nursery, as well as later while working for the Arnold Arboretum, and he noted in his collection notebooks that the species was \"very rare!\" This characterization stands in stark contrast to his observations of the common Japanese species. Hobbled Horticulture No matter the botanical interest of these species, it was another thing to bring them into more widespread cultivation. In 1889, Charles Sprague Sargent, the founding director of the Arnold Arboretum, wrote a short article about Viburnum lantanoides for Garden and Forest. He praised the species as \"one of the most beautiful plants of our flora,\" yet he noted that the hobblebush was \"impatient of cultivation\" and \"the most difficult of all our native shrubs to cultivate.\" At the Arnold Arboretum, only two early hobblebush accessions achieved long-term success. Both arrived as plants and perished in the mid-twentieth century. Repeated attempts to cultivate the species since then--via seeds as well as layers from the wild--have been in vain until recently. Seeds refused to germinate, and the plants rarely lived beyond a few years after planting out. However, several plants collected in the Adirondacks in 2008 have survived near a swampy spot adjacent the North Woods, known for a planting of corkwood (Leitneria floridana)--the same location where one of the two earlier successes had grown. The other three species have been equally slow to enter North American gardens. Sargent collected seeds of Viburnum furcatum from Japan in 1892, yet no records of successful germination (or planting) exist. It was not until early 1915 that two collections from Ernest Henry Wilson in northern Honshu made their way to the Arnold Arboretum. One collection originated as plants, the other as seeds, and the resulting accessions (numbers 17988 and 17989, respectively) were also sited near the Leitneria. The year those accessions arrived, Sargent noted that the species was \"as handsome a plant as the American species, and will probably prove equally difficult to manage.\" However, the seed-derived accession survived until 1997, occasionally dying to the ground and sending up suckers. The other accession (containing three plants) has persisted in the location for a century, putting on dramatic floral displays typically in late April to early May. Another seed-derived accession arrived in 1998. Perhaps because this species takes more readily to various propagation techniques, Viburnum furcatum seems to be the most widely cultivated of the four species, showing up in some forty-three public gardens in fourteen different countries according to Botanic Garden Conservation International's PlantSearch database. The American species is growing in twenty-seven gardens in ten countries, while V. nervosum and V. sympodiale are growing in only eleven and nine gardens, respectively. These numbers, while likely accurate in the ranking of each species' popularity (or amenability to cultivation), are likely underestimates, for the database requires individual gardens to self-report what they are growing. For instance, no gardens in Japan are noted as cultivating V. furcatum, and they likely are. The Morris Arboretum has two specimens of Viburnum nervosum, which were acquired as cuttings from a specialist nursery in 2010. The Arnold Arboretum has only attempted to cultivate the species twice--two seed collections in the early 2000s--and, in both cases, the seeds failed to germinate. Wilson had made several herbarium collections of V. nervosum (as V. cordifolium) in western Sichuan, where he observed it was \"common\" and formed \"thickets.\" Joseph Rock also collected the species in high plateaus of Yunnan and noted it growing as both a shrub and even a tree up to twentyfive feet in height. Yet, the earliest record of the plant's introduction to North America seems to be a 1927 accession donated to the United States Department of Agriculture by the Lloyd Botanic Garden in Darjiling, India. As for the final species, Tony encountered Viburnum sympodiale in September 2005, 22 Arnoldia 76\/4 ? May 2019 from the Royal Botanic Garden, Edinburgh, which rooted and survived on the grounds for eight years. Three seed acquisitions were never successful. Disjunct Reunions If the early work of plant explorers enabled discoveries pertaining to biogeography, like Gray's initial articulation of disjunct species, then the slow acquisition of other disjunct species (including Viburnum nervosum and V. sympodiale) makes us wonder what breakthroughs current and future plant exploration will enable. Aside from Viburnum furcatum, our expedition to Japan yielded a number of other collections of species that have familiar North American relatives. Among the most exciting was a collection of Stewartia: another genus with members that only occur in eastern North PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY; PEABODY.YALE.EDU. PHOTO BY PATRICK SWEENEY while collecting in Gansu Province, China. Although there were no fruits, he and the other explorers gathered leaf tissue and herbarium specimens for future molecular work, and longterm preservation, respectively. This experience caused another d?j? vu moment all over again, and the field notes for the collection (NACPEC05-033) acknowledge \"this species is very reminiscent of Viburnum furcatum or V. lantanoides.\" The team found it in a mesic, mixed deciduous and coniferous forest--much like the habitats preferred by the Japanese and American species--replete with three maple species and a hemlock (Tsuga chinensis), as well as littleleaf boxwood (Buxus sinica), rosy dipelta (Dipelta floribunda), Rosa davidii, and katsura (Cercidiphyllum japonicum). Despite four attempts, the Arnold Arboretum has only had this species in cultivation once: cuttings Like both the North American and Japanese Pseudotinus, Viburnum sympodiale grows in moist woodlands. Here it flowers within a roadside forest in Hunan, China. The young secondary forest included an overstory of pines (Pinus), along with planted larches (Larix). ANTHONY S. AIELLO ANTHONY S. AIELLO Viburnum 23 While in Japan, the authors and collaborators collected fifty-five unique species. Half of the genera, like Viburnum, co-occur in eastern Asia and eastern North America. At left, collaborators Tatsuhiko Shibano and Mineaki Aizawa (left and right) scan the canopy of a steeply sloped forest at the University of Tokyo Forestry Department Research Station, searching for fruits of Stewartia pseudocamellia. The striking bark of this species is show at right. America and eastern Asia. At the University of Tokyo Forestry Department Research Station, in Chichibu, west of Tokyo, we encountered a population of Japanese stewartia (S. pseudocamellia) perched on a steep hillside, the trunks elongated, stretching for light among an overstory of impressive beeches (Fagus japonica). This stewartia, known for its exfoliating bark, large white flowers, and rich autumn leaf colors, is among the most ornamental plants that have come to our gardens from Japan. We were excited to make this collection because, despite its common name, our collecting companion Todd Rounsaville confirmed that the only documented, wild collections of Japanese stewartia in the United States are of Korean origin. Our collection represents an infusion of novel Asian material into cultivation. We also collected a second species of Stewartia at the last location of the expedition, the Hokkaido University Forestry Station in Wakayama Prefecture on the Kii Peninsula, the southernmost tip of Honshu. This mountainous region is among the wettest areas of Japan, receiving upwards of 118 inches (300 centimeters) of annual precipitation, which we experienced during a continuous downpour--one of the wettest collecting days of our collective careers. We made collections of Stewartia monadelpha, aptly named the tall stewartia, which grew to about 40 feet (12 meters) tall, with trunk diameters of nearly 18 inches (45 centimeters)--showcasing the slick, orange-red bark. Of course, we made other collections of disjuncts on the trip, including green- or snakebarked maples (Acer micranthum and A. ANTHONY S. AIELLO ANTHONY S. AIELLO ANTHONY S. AIELLO 24 Arnoldia 76\/4 ? May 2019 In addition to collections of familiar genera such as Viburnum and Stewartia, the expedition yielded unusual species including wheel tree (Trochodendron aralioides), which have no disjunct counterparts. At top, Michael holds a specimen of Trochodendron, which collaborators Tatsuhiko Shibano and Mineaki Aizawa (left and right, bottom left) collected from a towering specimen along one of the many mountain torrents charged with the recent rains. Viburnum 25 tschonoskii) that are within the same section or clade as our familiar North American moosewood (A. pensylvanicum). We also couldn't help but notice the genera without disjunct representatives. On the rain-soaked day in Wakayama, we collected three species of Enkianthus (E. cernuus f. rubens, E. nudipes, and E. sikokianus), a genus in the heath family (Ericaceae), which is represented by six species in Japan, seven in China, but none in North America. It was remarkable to see three distinct species growing together on one mountainside. Earlier on the trip, we collected E. campanulatus and E. subsessilis, bringing our total to five of the six Japanese species. We also made two horticulturally and botanically interesting collections at Wakayama: wheel tree (Trochodendron aralioides) and Japanese umbrella pine (Sciadopitys verticillata), which both have a peculiar taxonomic standing. Wheel tree has only one other member in its family and order (Trochodendraceae and Trochodendrales, respectively), and the umbrella pine is the lone representative of its family (Sciadopityaceae). Importantly, through collection we've brought these species--long-lost cousins (as well as evolutionary orphans)--together in common gardens, as living plants, to observe them in cultivation. And like our pursuit of these four d?j? vu viburnums, the work as plant explorers continues. Gray used collections of herbarium specimens as inspiration to make daring hypotheses about biogeography. In that same vein, as plant explorers and curators we build collections to inspire future scientists to make new discoveries using not herbarium specimens and DNA samples (important as they may be) but living organisms. It is a longgame we play, however, for a century after the last species in the quartet was described, it has nearly evaded our cultivation. But, with dogged determination, hopefully we (or our successors) will achieve the perfect ensemble: a full quartet growing and performing together for audiences to enjoy and to study. Further Reading\/Bibliography Diels, L. 1901. Die Flora von Central-China. Botanishche Jahrb?cher f?r Systematik, Pflanzengeschichte und Pflanzengeographie (Vol. 29). Leipzig: Wilhelm Engelmann. Don, D. 1825. Prodromus Florae Nepalensis. London: J. Gale. Donoghue, M.J. 1981. Growth patterns in woody plants with examples from the genus Viburnum. Arnoldia, 41(1): 2?28. Edwards, E.J., D.S. Chatelet, L. Sack, and M.J. Donoghue. 2014. Leaf life span and the leaf economic spectrum in the context of whole plant architecture. Journal of Ecology, 102: 328?336. Ferguson, I.K. 1966. The genera of Caprifoliaceae in the southeastern United States. Journal of the Arnold Arboretum, 47: 42?43. Gray, A. 1859. Diagnostic Characters of New Species of Ph?nogamous Plants, Collected in Japan by Charles Wright, Botanist of the U. S. North Pacific Exploring Expedition. Memoirs of the American Academy of Arts and Sciences, New Series 6(2): 377?452. Mackenzie, K.K. 1927. A botanical riddle. Torreya, 27(5): 81?83. Marshall, H. 1785. Arbustrum Americanum: The American grove. Philadelphia: Joseph Crukshank. Michaux, A. 1803. Flora Boreali-Americana (Vol. 1). Paris and Strasbourg: Levrault. Maximowicz, C.J. 1880. Diagnoses plantarum Novarum Asiaticarum. St. Petersburg: Imperialis Academiae Scientiarum. Park, B., M. Sinnott-Armstrong, C. Schlutius, J.-C.P. Zuluaga, E.L. Spriggs, R.G. Simpson, E. Benavides, M.J. Landis, P.W. Sweeney, D.A.R. Eaton, and M.J. Donoghue. 2018. Sterile marginal flowers increase visitation and fruit set in the hobblebush (Viburnum lantanoides, Adoxaceae) at multiple spatial scales. Annals of Botany, 123: 381?390. Sargent, C.S. 1889. Viburnum lantanoides. Garden and Forest, 2(89): 531?532. Sargent, C.S. 21 June, 1915. Viburnums of Western Asia. Bulletin of Popular Information, 1(9): 33?35. Anthony S. Aiello is the Gayle E. Maloney Director of Horticulture and Curator at the Morris Arboretum of the University of Pennsylvania in Philadelphia. Michael S. Dosmann is keeper of the living collections at the Arnold Arboretum. The authors thank Michael Donoghue, Brian Park, and Patrick Sweeney for their conversations in the preparation of this article and use of images, as well as our collecting companions during our expedition to Honshu in 2018 for their friendship and collaboration. "},{"has_event_date":0,"type":"arnoldia","title":"In Bornean Rainforests: Exploring the Flora","article_sequence":3,"start_page":26,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25661","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d0608926.jpg","volume":76,"issue_number":4,"year":2019,"series":null,"season":null,"authors":"Ashton, Peter Shaw","article_content":"ASHTON, P. 2019. IN BORNEAN RAINFORESTS: EXPLORING THE FLORA. ARNOLDIA, 76(4): 26?39 In Bornean Rainforests: Exploring the Flora Peter Ashton A expeditionary qualifications had been on family holidays in France and Italy. I went to Brazil as an entomologist, and I returned with a keen interest in forest botany. John Corner, a professor of tropical botany at the University of Cambridge, encouraged my new enthusiasm, and he promised to pass on any leads about career opportunities for a budding field worker. Thus, my arrival in Brunei. We would be leaving for the Temburong River the following morning, Bill informed me. This river flows through the eastern portion of Brunei--which is divided into two noncontiguous sections--and once we departed from the capital city, familiar accommodations would be scarce. Bill scrutinized my appearance--athleticwear that, in retrospect, was more suited for camping in the English countryside--and he pointed me in the direction of a provisions store. \"You had better go and buy some clothes,\" he said. t twenty-three years old, I found myself on an open-aired launch, motoring from the island of Labuan, off the northern shore of Borneo, and entering the humid mouth of the Brunei River. The riverine settlements near Bandar Brunei, the capital, sprawled before me. Houses and other structures rose on stilts, interconnected with bridges and walkways that formed a vibrant and densely populated community, all above water. Boats crisscrossed in every direction. It was 1957, more than two decades before I would be appointed the sixth director of the Arnold Arboretum, and I was arriving in Brunei to work as the first official forest botanist for the Bruneian government. At the time, I could scarcely have predicted that this would mark the beginning of five years working under the auspices of Omar Ali Saifuddien III--the Sultan of Brunei--or, for that matter, that it would mark the beginning of a lifetime of research on the forests of northern Borneo. At Bandar Brunei (now known as Bandar Seri Begawan), I was welcomed by Bertram (Bill) Smythies, who held the title of state forest officer. Bill was the author of classic texts on the birds of Burma and Borneo. His father and grandfather had been botanists in the Indian Forest Service, and as he stood, awaiting my boat on the north side of the river, I could immediately see that his gaunt and sinewy physique had been shaped by his ceaseless fieldwork. During World War II, he had been protected behind Japanese lines, within Kachin longhouses, while he continued his research in northern Burma. I later learned from my local team members that he spoke their language (Iban) so fluently that, were he hidden from view next door, he would not be recognized as a foreigner. By comparison, I was exceptionally new to this world. My first and only prior field experience occurred in 1954, at age twenty, when I spent my freshman summer with two friends on the northern bank of the Amazon River, near Monte Alegre, Brazil. Before that, my only As a new forest botanist, I had been charged with gaining field information about the tree flora of a region slightly larger than Delaware, concentrating on the dominant tree family, Dipterocarpaceae. Dipterocarps are colossal monarchs of the paleotropical rainforest, dominating the overstory. Over the previous half century, they had become the leading source of general-utility timber, and they also provided quality hardwood for flooring and decks. Because dipterocarps were entering the regional export market, I was to provide preliminary information that would guide sustainable management for this timber. I had visited the herbarium at Royal Botanic Gardens, Kew, near London, to read up and see examples in the herbarium. But the specimens revealed a diversity of foliage that was unfamiliar and complex. My knowledge, at the start, was little more than rudimentary. Before my departure from Bandar Brunei, I learned that my office in town lacked botany books and that no tree flora for Borneo had been published. Moreover, the herbarium at the office consisted of a closet with a small col- B RU N E I B AY Bandar Seri Begawan SO U TH CH INA S EA mb Te Tutong ur o n gR iv e r Li mb a n Andulau Hills g R iv Kuala Belait Kuala Belalong er B RU N E I B RU N E I Bukit Retak Gunung Pagon Periok MAL AY SIA (Sarawak) B OR N E O 1 0 M I LE S 2 5 KI LO M E T E R S Ashton arrived at Bandar Brunei (now known as Bandar Seri Begawan, top left) in 1957. His research in the Bruneian forests between 1957 and 1960 focused on overstory species like this giant dipterocarp, Shorea longisperma (top right), and followed routes mapped in red. MAP BY PETER ASHTON, ARNOLD ARBORETUM, AND GIS COMMUNITY PHOTOS BY AUTHOR FROM ARNOLD ARBORETUM ARCHIVES UNLESS NOTED Peter Ashton in Brunei 27 28 Arnoldia 76\/4 ? May 2019 together more briefly). The Iban are members of an inland minority on Borneo, known collectively as the Dayak, which includes major groups that speak distinct languages. Later, an older and more experienced Iban collector named Sengelang anak Nantah joined my team. Given his considerable experience and wisdom, I called him Apai, meaning Dad. My field companions possessed an optimistic, energetic and ribald self-confidence. That first evening and night was both exciting and reassuring. My companions were endlessly helpful and friendly, although already detecting opportunities to pull my leg. The rush of the rapids, with the calls of the evening cicadas, frogs, and birds, was both exotic and inviting. I was already familiar, from student days, with the customary method of collecting botanical specimens and inserting them in wooden presses to dry, but the special requirements of doing so in dense forest under a climate of intense humidity and heat was challenging. The aim TIMOTHY CHARLES WHITMORE lection of specimens, mostly from the 1930s. I needed help, so Geoffrey (Geoff) Wood, a forest botanist from adjacent Sabah--now a state in Malaysia--came down to be my botanical tutor for the first three months. Geoff had hurt his leg in a field-hockey game before the expedition, causing his ankle to swell painfully, but he was nonetheless a tireless worker, and his field experience proved invaluable. It took us three days to reach Kuala Belalong, the fork where the Temburong River meets the white waters of the Belalong River, deep between hills that rise nearly 2,000 feet (600 meters), enshrouded in mighty emergent trees. I had learned much on the way up: I now knew that my life would revolve around an outboarddriven prahu (dugout canoe). I knew that I needed to learn the lingua franca Malay, and fast, and that Iban would be useful as well, given that I would spend day and night with a team of Iban lads near my own age: Ladi anak Bikas, Asah anak Unyong, and Naban (whose paternal name I do not recall, because we worked The field team (from left) included Asah, Naban, Ashton (wearing his indispensable leech socks), and Ladi. Peter Ashton in Brunei 29 Ladi demonstrates how diagnostic notches are made in the trunk of a large dipterocarp species, Shorea slootenii, using a parang. At right, a leaf is shown with the notched trunk of Dipterocarpus conformis. was to collect eight quality vouchers of every tree in flower or fruit, to send to regional herbaria, as well as to leading herbaria worldwide, including at Leiden University, Kew, and yes, the Arnold Arboretum--my first introduction! As a newcomer, the challenge of identifying giant trees, towering more than 200 feet (60 meters) overhead, quickly introduced me to the parallel world of Iban forest botany--a world very independent of the formal plant science of a western university. Flowers provide the best means to identify the botanical family, using methods devised by Linnaeus, but discovering fallen petals (or fruit) was rare. Canopy species only start to flower once their crown is in the sun. Few Bornean species flower annually, and some go many years without. If they do flower, however, it is most likely between the two wet monsoons, in and around April, when sunny periods of a few days are frequent in most years. So this was always the time to plan major collecting expeditions, and it was, in fact, around this time that our trip commenced. As I began to grasp a little of the Iban language, I started to ask Asah for help. Asah's approach was, first, to note the overall bark appearance, observing its color and whether it was smooth, flaky, fissured, or otherwise. He would then slice into the bark. He would smell the wood, look for sap or latex, and note the overall color of the inner bark and sapwood, as well as the presence of radiating pale lines (known as medullary rays). For this purpose, we all wore a parang at our waist--a short, sharp blade resembling a billhook without a bill, crafted by a blacksmith from a truck spring. (Parangs were also used for cutting paths and much more.) These wood and bark character- 30 Arnoldia 76\/4 ? May 2019 near Los Angeles--Santa Barbara, Laguna Beach, Ventura--from where they had been exported as packing material. The packed specimens were then doused with a mixture of 90 percent ethyl alcohol, fortified with formalin and mercuric chloride powder, which helped to preserve the specimens until they could be dried in presses at the herbarium and dispatched or mounted. istics provided enough information to home in on the family and often the genus, for which Asah and the other collectors would often know an Iban name. Eventually, I learned that nearly all species could be identified on the basis of their leaf and petiole morphology, at times with the added assistance of trunk characteristics or a fallen twig fragment, without requiring flowers or fruit. Bill and I would bring samples back to the camp for Geoff to examine. In addition to providing a loose identification for the specimen, he could tell us whether something had already been well-documented with other herbarium collections. With this guidance, we knew what to collect. At this point, the fun really began. Asah, Ladi, and Naban were extraordinary tree climbers. (We never needed to fell a tree to collect specimens.) The men would clasp the trunk in a crouched position, using their arms and the soles of their feet, and then stretch up to embrace the trunk again. The trunk needed to be sufficiently slender to clasp, so if the target tree was too large, the climber sought out a smaller tree nearby. He would climb this neighboring tree until the stem began to bend, and then, using his weight to swing the crown of his tree back and forth, he would lean towards the next tree and hook a long, forked pole (known as a penyulok) around a branch. He could then tie the lower end of the penyulok in place, bridging the two trees. By hauling up a new penyulok on rattan cord, the procedure could be repeated until the main branches of the emergent tree were reached. The rest depended on confidence, agility, and iron will! The twigs, once dropped from on high, would be tied in bunches and placed in large plastic bags familiar to field botanists. I initially learned the Schweinf?rth method of preparing specimens, which was developed for wet, hot climates in the region by Dutch botanists. The method requires rustproof metal containers, which were made by local tinsmiths. These were designed with a cross-section large enough to fit specimens, which were entered between sheets of newspaper. It was both surreal and entertaining, in the gloaming of a rainforest evening, to spend moments reading the goingson in the local newspapers from seaside locales Bornean forests are not dangerous places, so long as care and discretion are the rule. Beyond villages and longhouses, mosquitoes and sandflies (no-see-ums) are confined to the riverbanks. Leeches are a perpetual pest, but they can be excluded by wearing gabardine kneelength stockings. Dangerous animals are few, although I did manage, a few months later, to step on a pit viper, after which Ladi carried me on his back for two days across high hills to the boats. Nevertheless, inland roads were scarce, and ascending white-water rivers presented a perpetual hazard. Moreover, the dense forests of undulating lowlands created conditions where one could easily get lost, especially on cloudy days, when the sun provided no bearing. Scarcely two months into my time in Brunei, however, I learned a sobering lesson about backcountry perils. The mixture of chemicals used for the Schweinf?rth were dangerous, even to the touch. I would later search for an alternative, but tragically, while we were breaking camp in the Andulau hills--near the coast, west of Bandar Brunei--a five-gallon drum of ethyl alcohol, standing too close to a camp fire, exploded on Geoff. One of our Iban assistants ran for help and managed to secure an Australian roadbuilder with a short-wheelbase Land Rover, in which we carried Geoff, fifteen miles along the beach at high tide, to an oil-field hospital. He did not recover. The experience was traumatic, and I recall riding with Geoff's climbers--Kadazan men from Sabah--to the little graveyard in Kuala Belait, on the western coast of Brunei. All of us wept. The Kadazan collectors returned home, while I would return to the field. Geoff, who I had not known before, was patient and immensely knowledgeable. We had shared long evenings of relaxed banter under canvas, and he could not have been a better instructor in the art and science of tropical forest botany. SPECIMENS FROM HARVARD UNIVERSITY HERBARIA GEOFFREY WOOD Peter Ashton in Brunei 31 Collectors on a later expedition in Sarawak rest while setting up camp (top left). Schweinf?rth tins--used for preserving herbarium specimens--are stacked in the foreground. Asah and Naban are photographed climbing Shorea curtisii (top right). Asah has reached the basal branch of the crown; Naban is positioned about halfway up the trunk, where a small neighboring tree has been bridged with the Shorea; and Ashton stands at ground level. Ashton sent specimens to the Herbarium of the Arnold Arboretum, including Shorea amplexicaulis (fruits at right) and Shorea laxa (at left)--both collected in 1958. 32 Arnoldia 76\/4 ? May 2019 The tidal rivers of Brunei are inhabited by taxonomically diverse mangrove species like Avicennia alba (top), Heritiera globosa (along with nipa palm, Nypa fruticans, bottom left) and Rhizophora apiculata (bottom right). Notice the unusual seedlings of the Rhizophora, which germinate and produce long drooping stems while the seeds are still attached to the parent plant. Peter Ashton in Brunei 33 Brunei is a small place, and most collecting trips took only a few days, although even those entailed slow travel up small rivers, with a nightly welcome in longhouses or forest camping beneath waterproof flysheets. Once a year, throughout most of my five years in Brunei, I took advantage of expected drier times in April to mount a major expedition. (I did the same in other parts of Borneo over the subsequent five years, amounting to six expeditions in all.) The first, undertaken when I had been in Brunei just over one year, tried my organizational and leadership skills and proved among the most eventful. Not only had my familiarity with the trees grown substantially over the previous year, but I had also learned sufficient Iban to get the gist of the endless exchanges of robust humor in the boats as we travelled. My knowledge of Malay was growing as well. My first long backcountry expedition targeted Brunei's highest mountain, Gunung Pagon Periok, which is a steep sandstone slab that rises over 6,000 feet (1,850 meters) at the cascading headwaters of the Temburong River in the eastern portion of the country. We would access the peak via a montane ridge in nearby Sarawak. With that elevation, I suspected that it should be possible to collect within all equatorial forest types, which are differentiated according to altitude. Pagon is visible on the horizon from Bandar Brunei, the capital city, but although it is hardly more than fifty miles distant, I knew from aerial surveys and maps that it would take many days to reach. The length of time we could spend hiking in the rainforest depended on the number of field assistants required to haul equipment and food, especially rice. The basic backpack is a selabit, woven from rattan. Its base fits snuggly in the small of the back, while the top reaches the crown of the head. For a two-month trip that required overland travel, we would need about fifteen field assistants--recruited along the way--in addition to the Iban climbers. Before heading towards Pagon, a day was spent shopping for sufficient food to last six to eight weeks: rice, cooking oil, salt fish, little red onions, chilies, and all-important matches and kerosene. I departed Bandar Brunei with Apai and Ladi, winding through vast mangrove swamps in an outboard prahu. We crossed the Bay of Brunei and entered the mouth of the Limbang River, whose valley separates the two halves of Brunei. This swamp, bordering the whole extent of the bay, harbors an exceptionally diverse flora, as well as a rich fauna including the notorious long-nosed, white-faced proboscis monkey (Nasalis larvatus). We then continued up the winding river through fertile floodplain farmland, to the large and elegant Iban longhouse of Tanah Merah. We were welcomed with a ceremonial bowl of tuak (rice wine), which was followed with a dinner of rice and chicken, loaded with plenty of chilies, and traditional dancing in the evening. I was cajoled into attempting a sword dance, which caused much hilarity. We managed to attract six enthusiastic assistants and a second outboard prahu, and we set off early again upriver. On a daily wage equivalent to my climbers (but lacking their field allowance), these recruits soon became socially part of our team. About midday, the current quickened, the occasional rapid had to be maneuvered, and the hills began to close in on both sides. We were now in the country of the Murut, a different ethnic nation from the Iban, but with a long-standing friendly relationship. When we stopped in a village, our welcome was subdued, as many were away, tending to their paddy farms. We spent our second night in one of their longhouses, which smelled strongly of rancid tuak--a sign that harvest festivities were over. We did succeed, however, in gaining another motorized prahu and a second contingent of young woodsmen so that our party, now fifteen, was complete. At noon on the third day, we reached Nanga Medamit, the tributary that drains the western flanks of Pagon. We pulled the prahus up the hillslope above the river, above highest flood levels, packed our equipment and food into selabits, and ascended the steep slopes to the ridge where we set up camp for the night. This gave me my first opportunity to chat with the Muruts, who were more reserved than my boisterous Iban companions. I had planned to use the expedition to start recording medicinal plant uses. One Murut seemed quite knowl- 34 Arnoldia 76\/4 ? May 2019 On the way to Gunung Pagon Periok, Ashton encountered a lower montane forest, known as kerangas, for the first time. During subsequent fieldwork in kerangas forests, Ashton photographed lipstick palm (Cyrtostachys renda, left), named for vibrant red leaf sheaths that do not appear in the photograph, and Borneo kauri (Agathis borneensis, right) a member of the coniferous family Araucariaceae. A Malay collector named Karim is pictured. edgeable about plants, but, on seeking his experience, he answered that, although his people were unsurpassed in their knowledge of plants that increase the hunting and sniffing skills of their dogs, all that they knew for humans concerned poisons. The following day, we successfully shot a fat boar, the Bornean bearded pig (Sus barbatus). Some was consumed on capture but most was boiled with salt and stored for the weeks to come. Time being limited, we decided to collect little until our base camp was achieved. This gave us the opportunity to observe changes in forest structure and flora as altitude increased. We followed the ridge upward, trading the sounds of the water for the occasional ghastly cackle of the helmeted hornbill (Rhinoplax vigil)--a call that now brings back happy memories. We came upon a large water-filled wallow that, I was assured, had been made by a rhinoceros-- the little two-horned species (Dicerorhinus sumatrensis), also found in Sumatra. In late afternoon, we scaled a small peak, about 2,100 feet (650 meters) high, where I detected an upper dipterocarp forest for the first time, just below the cloud base. These trees were shorter of stature with unfamiliar species including a dipterocarp that later proved new to science: Shorea flaviflora. The next morning, we started an ascent to about 2,800 feet (850 meters), which led us into a lower-statured woodland, lacking emergent trees but with a profusion of trees in the oak family (Castanopsis, Lithocarpus). This was my first experience of a lower montane forest. It differed from textbook descriptions in its open canopy and dense understory of polesized trees. I later learned this was a forest type Peter Ashton in Brunei 35 flat densely wooded southern slope facing us. We spent three weeks there, exploring, collecting, and setting out one-acre plots (we had not become metric in those days), which we fully censused. We scaled the perilously narrow summit ridge, which bore a short shrubby thicket, with little in flower. But the forest around the camp yielded exciting new discoveries including, amazingly, a new dipterocarp, which I would name Shorea monticola. After nightfall one rainy day, we saw a curious pale globe of light, less than a meter in diameter, moving slowly downhill through the distant trees, eventually disappearing. Could it have been a form of ball lightning? It did not last long, and my campmates were unperturbed, casually explaining it to be a benign forest spirit, going about its business. known as kerangas, which is widespread on Bornean mountains. The going got steeper as we approached a high spur, where, at 5,700 feet (1,750 meters), the trees were hardly more than head height. Everything, including the contorted branches of the trees, became so carpeted in moss that their dwarf crowns could hardly be distinguished from the ground, giving purchase to spectacular orchids and occasional rhododendrons (Rhododendron sect. Vireya). This was the upper montane forest, daily immersed in fog, dripping, and so unnervingly silent that our voices hardly carried more than a few yards. We sought out a campsite while Apai and Ladi pressed on, cutting through the vegetation with parangs. The only visible way through was a tunnel made by wild boar. So thick were the moss tussocks that we had to stay two more nights while a passage was cut to squeeze through with the baggage. We set up our permanent camp in an ecotone between lower montane kerangas and upper montane thicket, where we could see the base of Pagon's imposing sandstone slab, with its After five weeks in the forest, rice was beginning to run low. We had to retain enough supplies for our return when the weight of the food would be replaced by that of the full cases of specimens. Because we had accessed the mountain The collectors viewed the sandstone slab of Gunung Pagon Periok, with its steep slopes carpeted with upper montane forests, from their ridge of approach in early morning. 36 Arnoldia 76\/4 ? May 2019 from the Limbang River--which runs through Sarawak rather than Brunei--I discussed return options with Apai and our other teammates and was attracted to the idea, if possible, of continuing southeast with a few of our most energetic team members to access the Temburong River headwaters, which we would follow until the stream became navigable. This area had been explored by an oil geologist thirty years earlier, but it was otherwise unknown to western researchers. Apai confidently assured me that, upon reaching the Temburong, a bark raft could be constructed. So the main party returned the way we had come, carrying most of the baggage, while Apai, Ladi, one Murut assistant, and myself bid them farewell and set off, lightly equipped with five days of rice and other basic foodstuffs. That first day we made a steep descent of the northern slope of Gunung Pagon Periok to a ridge that connected to a lesser peak, known as Bukit Retak. Apai suggested that we should scale this peak to search out our route and possibly spend the night on the summit where we could usefully commune with mountain spirits. I succumbed to the idea, so we spent a soggy fog-enshrouded night beside a summit dewpond--but to no avail. We camped in an open valley the next evening and spent the following day scaling Bukit Lesong, the northernmost of Brunei's mountains, where I found another dipterocarp of unfamiliar leaf shape (which I would later know as a species of Vatica). Because we were traveling light, we had no means to collect specimens. On the following day--our third since leaving the others--we descended for five hours and came on the main Temburong stream, but it was flowing over massive boulders and was far too steep to attempt navigation. We followed it until evening, and at daybreak the next day, we searched for a tree with suitable bark for a raft. No such tree was to be found, so we felled the next best. The result was riverworthy but pliable, resembling a giant banana skin. We had not gone far that morning when we heard a After Ashton and his team finished their fieldwork near Gunung Pagon Periok, they descended towards the headwaters of the Temburong River, photographed on a later trip near Kuala Belalong. Peter Ashton in Brunei 37 Within the forests of the Temburong District--the easternmost enclave of Brunei--Ashton observed large overstory species like Anisoptera costata (left), which is a member of the dipterocarp family (Dipterocarpaceae), and Tristaniopsis whiteana (right), a member of the myrtle family (Myrtaceae). An Iban collector named Mujah is pictured. late. We leapt into the torrent, while our Murut assistant attempted to haul in the raft by its attached cord. All to no avail. The raft turned, bent in two, and flipped over the fall, and within it our clothes (including our shoes!), our remaining food, and our parangs--everything. The three of us crept to the edge of the fall and looked down. All that came up, turning in the whirlpool, was Apai's bamboo cigarette container which, after rescue, revealed five vital matches. massive clap of thunder over the mountains to the south and soon noticed a rise in the waters. We leapt to shore, dragging our craft as high up the steep hillslope as we could. Within little more than a minute, the waters had gone up fifteen feet, swirling, carrying whole trees, the rocky bottom shaking and rumbling. No further travel was possible until the following day, when the flood had somewhat subsided but the current was still strong. We proceeded with caution. Ladi and I had ascended the Temburong some months earlier, and we were stopped by a two-meter waterfall called Wong Uan, which was practically impossible to portage or descend. Below it, a cataract known as Gerugu Rimau raged between the cliffs. We knew we were getting close. After little more than one hour on the river, we rounded a bend, and Ladi and I recognized the terrain--only too Our only option was now to find the nearest Iban longhouse, several days walk downstream. We rested, and the next day--the fifth since we left the main party--was a disaster: showery, cloudy, with no clear view. Walking barefoot in the rainforest proved easier than I had imagined, but in late afternoon, we began to identify bent twigs with which we had marked our morning 38 Arnoldia 76\/4 ? May 2019 We waited and waited. Three days passed. Ladi became increasingly concerned and eager to find a floating log himself, but I discouraged him. Then, on the fourth day, the distant noise of an engine could be heard, coming, then going. That's an airplane, Ladi said in sad conviction, but I was more confident. Sure enough, after a long wait, around the river bend came a prahu manned by my friend Penghulu Gimang (an upriver chief), with his son Jah. They had been alerted by Apai and his Murut companion, and they brought a feast: salt fish, over which they poured condensed milk. Never, ever, have I tasted anything so delicious! We descended the river, stopped at Gimang's longhouse, and then proceeded down to the estuary and across the bay to the capital. On the way, we noticing occasional shirts, towels, and other flotsam high in the branches of the overarching trees. trail. We had turned in a complete loop. We set up a shelter by snapping off leaves from a fan palm (Licuala), and we found dipterocarp resin within a hollow tree, which, with some tinder, allowed us to start a fire. We spent that night toe-to-toe around the flame. The following morning, the sun revealed clear blue skies, so Ladi, climbing a tree, was able to discern the best way forward, bypassing a major turn of the river down to its confluence with a stream called Nanga Temburong Machang. By this time, Ladi was getting exhausted. We crossed the river, which had begun to subside, and Apai and our Murut assistant left us, floating on a log down the rapids in search of help. Ladi and I lit a fire with the last matches. We had plenty of water to drink, but food was sparse: some young turtles, which were exceedingly chewy after we cooked them in their shells, and the sweet and familiar fermenting pulp from the fallen fruits of a giant leguminous tree. We even took to eating clay to fill our stomachs, which turned them to cement. This initial immersion in Bornean forest botany lasted twenty-six months. During that period, Ladi and Asah became my friends for life. (We Wong Uan, a waterfall on the Temburong River, stranded Ashton's team without supplies. It was photographed here at more placid conditions in 1958. Peter Ashton in Brunei 39 Ashton's fieldwork documented 151 dipterocarp species in Brunei, including species like Dipterocarpus lowii (left), and Shorea rubella (right), which are both considered critically endangered by the International Union for Conservation of Nature. still communicate from time to time through a Malaysian friend.) We made nearly four thousand collections from which Dutch colleagues at Leiden University would assign over seven hundred scientific names. This was enough for Hasan bin Pukol--curator of our new herbarium in the attic of a local cinema--and I to publish a checklist of trees, which included local names. I also gained enough knowledge of the dipterocarps to publish a manual describing Brunei's 151 species, of which 33 were new and formally named for the first time (only 3 more have subsequently been discovered). This information would prove essential for future sustainable management and conservation efforts, and, on a personal front, these months in the field provided materials that would become the basis for my doctoral dissertation and a career beyond, including as the director of the Arnold Arboretum. Along the way, I had come to recognize the floristic and ecological patterns in the forest, and a newcomer--a neophyte, really--had found a way of life. My paleotropical education had officially begun. Peter Ashton is Harvard University Bullard Professor Emeritus and was director of the Arnold Arboretum from 1978 to 1987. Among many career honors, his research on tropical forests was recognized with the prestigious Japan Prize in 2007. He and his wife, Mary, live in Somerset, England. The map in this article was created using Esri, USGS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, Intermap and the GIS user community, based on an illustration that originally appeared in Ashton, P. 1964. A Manual of the Dipterocarp Trees of Brunei State and of Sarawak. Oxford: Oxford University Press. "},{"has_event_date":0,"type":"arnoldia","title":"The Old Timer","article_sequence":4,"start_page":40,"end_page":41,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25663","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d070a36f.jpg","volume":76,"issue_number":4,"year":2019,"series":null,"season":null,"authors":"Mayer, Bob","article_content":"MAYER, B. 2019. THE OLDER TIMER. ARNOLDIA, 76(4): 40 The Old Timer Bob Mayer M ore than ten years ago, I photographed an odd-looking tree growing in front of the Roxbury puddingstone outcropping on Valley Road. It was early May, and the tree was in bloom--sort of. One of the main branches was barren. It looked like the tree had been mistakenly passed over by the crews clearing away the dead and dying after a long winter. When I photographed the tree two years later, in 2009, the horticultural staff had visited, not to remove the tree but to prune it vigorously. The barren branch was gone, and the tree resembled a warty, one-eyed beast, sprouting feeble arms. I checked the metal tag: a Yoshino cherry (Prunus ? yedoensis forma perpendens, accession 22542*A). It was accessioned in 1925, a dozen years before my own birth date. Perhaps because of that, I developed an attachment to this aging--indeed ancient by cherry standards--tree despite is ungainly appearance, and I occasionally grabbed other images of the \"old timer,\" as I nicknamed it. While the record label suggests this accession was grown from seed sent from the Imperial Botanical Garden in Tokyo, Japan, I learned that an important intermediate step was involved. The tree was grown as a seedling from another Yoshino cherry (accession 5351*A), which arrived from the Imperial Botanical Garden (now known as the Koishikawa Botanical Garden) in 1902. The original tree grew near the Forest Hills Gate, and Charles Sprague Sargent often commented on the pink and white flowers. Even though the buds were regularly nicked by spring frosts, Sargent esteemed the hybrid as \"one of the handsomest\" cherries from Japan. According to Donald Wyman--a long-time horticulturist at the Arboretum--the Arboretum's original tree represented the first introduction of the Yoshino cherry into America. This predated a more famous gift of the hybrid (along with other Japanese cherries) to the city of Washington, DC, in 1912, where it formed the basis of the famous planting that clouds the Tidal Basin with evanescent blossoms each spring. This lineage is especially significant given that, when Ernest Henry Wilson visited Japan in 1914 and 1915, he reported that fortyyear-old trees at the Imperial Botanical Garden were the oldest known representatives of this hybrid (which is now considered a complex cross between Prunus speciosa and P. subhirtella) and that the original taxonomic description had been based upon them. Wilson observed the old Yoshino cherries flowering at the Imperial Botanical Garden, with benches where visitors could sit beneath the outstretched branches. Despite the recent scientific recognition of the hybrid, Wilson described its omnipresence throughout Tokyo. \"This is the Cherry so generally planted in the parks, temple grounds, cemeteries and streets,\" he wrote in The Cherries of Japan, published in 1916.\"Its flowers herald an annual national holiday decreed by the Emperor. In all over fifty thousand trees of this species are growing in the precincts of the city.\" This celebration, known as hanami, is still enormously popular in Japan, and it is premised on appreciating ephemerality--a celebration of fleeting beauty. After discovering the significant background of the old timer, I returned recently for another look. It seemed taller and statelier, now that I had uncovered its history. Horticultural care during my decade of observations had maintained--seemingly even resurrected--this old tree, which looked even healthier now than when I first encountered it. If the spring flowers symbolize the swift passage of the seasons, then the knobby form of this tree seems to extend this metaphor even further, embodying the passage of years. I'm confident the tree will survive much longer than this humbled observer. Bob Mayer has been birding, photographing, and volunteering as a docent and field study guide at the Arnold Arboretum since 2002. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23465","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14ebb6e.jpg","title":"2019-76-4","volume":76,"issue_number":4,"year":2019,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Hybrid Mystique","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25659","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d0608528.jpg","volume":76,"issue_number":3,"year":2019,"series":null,"season":null,"authors":"Grossman, Jake J.","article_content":"GROSSMAN, J. J. 2019. THE HYBRID MYSTIQUE. ARNOLDIA, 76(3): 2?13 The Hybrid Mystique Jake J. Grossman H ybrids, the results of successful breeding between two parents of different species, occur frequently in nature, yet are perhaps most familiar to us when they result from human intervention. We encounter in our intentionally cultivated hybrids the utility of the mule, the stateliness of the London plane tree, and the sensuous smells and tastes of myriad vegetables and fruits, including broccolini and the tangelo. These remarkable examples are of our own making, but hybridization between closely related species is perhaps the norm rather than the exception in nature. Though hybrid offspring are sometimes sterile and can be visually distinct from their parents (like mules), they are just as likely to be fertile and to pass unnoticed by us. These cryptic hybrids, diagnosable only through genetic testing, breed with each other or with individuals of their parent species (in a process called backcrossing), giving rise to new hybrid progeny. Over generations, such interbreeding consolidates novel hybrid traits, sometimes leading to the formation of new species. Because what counts as a species is, after all, merely conventional, it could be said that we humans, the descendants of interbreeding between Neanderthals and early Homo sapiens, are just as much hybrids as the most luscious of tangelos. Perhaps foremost among the natural world's \"hopeful monsters\"--a term that geneticist Richard Goldschmidt coined in 1940 for evolutionary transformations that occur through rare but large-scale mutation--hybrids often captivate and delight human observers.1 Yet hybridization does not always precipitate the formation of a new species. In natural populations, hybrids are frequently formed, only to be subsumed, through backcrossing, into their parental stock. This process--called introgression--results in the enrichment of the gene pool of the predominant species with genetic material from close relatives. So, in the case of oaks--described as particularly problematic for the biological species concept due to their wanton tendency to hybridize (Burger, 1975)-- we might say that there is evidence of red oak (Quercus rubra) introgression into a stand of northern pin oaks (Q. ellipsoidalis). These pin oaks will still be pin oaks, but perhaps with some hidden genetic diversity and leaves or bark that look, well, just a little bit different. The ubiquity of such situations has led biologists to formulate the idea of hybrid complexes or zones: sets of species or populations in which rampant interbreeding has produced a messy gradient of similar organisms, rather than discrete sets. Our cultivated citruses represent one such complex, in which ten progenitor species in southeast Asia and Australia have given rise, through hybridization, to dozens of domesticated taxa (Wu et al., 2018). And so, when I began my own foray into the world of hybrid aspens (Populus), I risked wading into a thicket of biological questions that could have been difficult or impossible to resolve. Fortunately, I was a first-year graduate student, a neophyte far more optimistic than I am now when it comes to tackling a new project. What follows is a story of a journey through which a team of ecologists and evolutionary biologists, myself included, tried to track down the truth about a putative hybrid. To do so, we traveled throughout the Midwest and dug deep into the natural history of the Niobrara River Valley, a relictual ecoregion left behind by the retreat of glaciers at the end of the last Ice Age. A biotic crossroads: The Niobrara River Valley Heading into the Nebraska Sandhill region along the state's border with South Dakota, new visitors might be surprised to plunge from cornfields and pastures stretching as far as the eye can see into forested canyons hugging a cool, inviting river. Originating on the eastern edge of Wyoming, the Niobrara River runs from west to east across the northern quarter of Nebraska NATIONAL PARK SERVICE NATIONAL PARK SERVICE Smith Falls Aspen 3 Smith Falls aspens (Populus ? smithii, right) along the Niobrara River, in northern Nebraska, are natural hybrids of two common North American aspen species. It shares habitat with northern species like paper birch (Betula papyrifera) and western species like ponderosa pine (Pinus ponderosa), both shown at left. before being subsumed in the Missouri River in the northeastern corner of the state. The canyons found between the river's banks and the surrounding matrix of arid short- and mixed-grass prairie constitute the Niobrara River Valley. For an eager observer, this valley presents more than just a respite from Nebraska's hot, dry uplands. Rather, as Midwestern botanist Charles Bessey first observed in 1887, the Niobrara River Valley is a \"meeting-place for two floras,\" a unique location in North America in which East and West comingle. Born and raised in Arizona, I had spent about five years living in the Midwest by the time I first visited the valley. When I arrived by car from St. Paul, Minnesota, where I had just begun my doctoral studies as a plant ecologist, it felt like I was seeing old friends again after a long absence. Most noticeably, stands of ponderosa pine (Pinus ponderosa), a decidedly western species, greet visitors to the Niobrara. These pines are among over a dozen western vascular plant species whose distributions extend all the way into Nebraska, following the biotic eastwest highway formed by the Niobrara River. A second look confirmed that this traffic moved in both directions: bur oaks (Quercus macrocarpa) and silver maples (Acer saccharinum) made unusual appearances for species that generally cannot be found in great abundance further west than the meeting of the prairies and forests in Minnesota and Iowa. It also became clear that the cooler, north-facing slopes of the valley, in particular, offered suitable habitat for species generally found further north, including cosmopolitan but drought-intolerant paper birch (Betula papyrifera). The same pattern-- a confluence of biota typical of the montane West, the deciduous forests of the East, and the boreal forests of the North--holds for herbaceous plants, insects, and vertebrates as well (Kaul et al., 1988). NATIONAL PARK SERVICE 4 Arnoldia 76\/3 ? February 2019 The hills and canyons of the Niobrara River Valley have provided shared habitat for an unusual combination of eastern and western species--plants and animals, alike. Of aspens, poplars, popples, and cottonwoods Any exhaustive flora of the Niobrara is bound to mention aspens, meaning species of the genus Populus. These trees go by a plethora of common names: aspens, poplars, popples, and cottonwoods--names which do not neatly map on to the current phylogenetic characterization of six sections within the genus (Hamzeh and Dayanandan, 2004). In North America, native species from section Populus, and some from Tacamahaca, are referred to as \"aspens,\" \"poplars,\" or \"popples.\" These include the ubiquitous quaking aspen (P. tremuloides), its close relative bigtooth aspen (P. grandidentata), and cold-tolerant basalm poplar (P. balsamifera). The species we know as \"cottonwoods\" are restricted to Aigieros and also to Tacamahaca, and these include, most famously, eastern cottonwood (P. deltoides, one of the largest trees east of the Mississippi), Fremont cottonwood out west (P. fremontii), and northwestern black cottonwood (P. trichocarpa), the first tree species to have its genome sequenced. Most species in the genus share traits with each other and with other members of the willow family (Salicaceae): they are dioecious, meaning that male and female flowers are borne on separate trees and have simple leaves and relatively short lifespans (often less than one hundred years). Many species have circumboreal distributions; they are generally cold tolerant, but vulnerable to hot and dry conditions. Aspens writ large have captured the imagination of botanists and the general public alike by virtue of their propensity for perpetuation through vegetative means. Most importantly, this means that a given aspen tree can, regardless of sex, produce new, genetically identical clones of itself. These new stems, often called \"suckers,\" emerge from rhizomes, underground stems that spread in parallel to the ARNOLD ARBORETUM, US FOREST SERVICE, AND GIS COMMUNITY Smith Falls Aspen 5 The distributions of quaking aspen (Populus tremuloides, shown in gray) and bigtooth aspen (P. grandidentata, shown in green) present a curious question: how could a hybrid between the two species occur in northern Nebraska, where one parent is exceptionally rare and the other is completely missing? soil surface. When a rhizome's buds encounter moist, warm conditions, a small shoot heads upward, sprouting leaves and emerging from the soil as though a seed had germinated in that exact spot. But these suckers grow faster than a typical seedling ever could, drawing on resources from their parental plant and bypassing seedling-hood in a mad dash for growth. These clonal offspring can, over time, become separated from their parents through the decay of the rhizome, but they remain genetic clones, such that what often appears to be a stand of aspen trees is, in reality, a single individual, connected, to varying extents, underneath the soil. The mystery of the Smith Falls aspens Aspen stands are vanishingly rare, if not altogether absent, in the Great Plains. Quaking aspens (Populus tremuloides) are nearly the sole representatives in the region but are restricted to a few tiny islands, usually growing in dense clumps around wetlands. These stands are relics, groups of trees left behind as the global climate warmed over the last ten thousand years. At the end of the Pleistocene Ice Age, aspen species, as with other cold-hardy trees, were probably quite common across this massive inland plain (Wright et al., 1985), but their ranges retracted as the climate became more arid. Stands of quaking aspen in central Canada and in the biotic highways of the Niobrara River Valley provide the only linkages between eastern and western populations for that species. My PhD advisor had secured funding from the National Parks Service--the Niobrara is a National Scenic River--to study one such stand, a rare, and therefore locally famous, collection of aspens centered around Smith Falls State Park, near Valentine, Nebraska. In 2013, we headed out for the first time to see these trees for ourselves. NATIONAL PARK SERVICE PAUL WRAY, IOWA STATE UNIVERSITY, BUGWOOD.ORG BECCA MACDONALD, SAULT COLLEGE, BUGWOOD.ORG 6 Arnoldia 76\/3 ? February 2019 Many morphological characteristics for the Smith Falls aspens (below) show intermediate traits between the parent species, including the shape and number of teeth on the leaf margins and the degree of leaf and bud pubescence. Quaking aspen (Populus tremuloides) is shown at top left, bigtooth aspen (P. grandidentata) at top right. Smith Falls Aspen 7 When we began, it was immediately clear that the aspens at Smith Falls bear a great resemblance to quaking aspens. They have lovely white stems and dark green, heart-shaped, finely toothed leaves. They grow in clumped stands, indicating spread by rhizomes, with no singletons off on their own. We found these trees growing in a string of ten stands extending along the Niobrara River Valley, from Nature Conservancy holdings in the east to private property in the west. The stands in between these two locations are one of the crown jewels of Smith Falls State Park and are well known to its many visitors, who also come to raft the Niobrara River, camp out along its banks, and see the eponymous waterfalls. The aspen stands are confined to the cool, north-facing banks of the river, and they're undeniably beautiful trees. But, as aspen fanatics, we agreed with what we'd already heard about them: they seemed somehow different than the quaking aspens so familiar to us in Minnesota. Their bark, though light-colored, seemed rather green, and their leaves were rather large, with fewer and larger teeth, compared to a typical quaking aspen. Indeed, these trees had long been held by local botanists and natural historians to be hybrids between locally rare quaking aspen and bigtooth aspen--a species whose eastern range edge is currently estimated to fall around 375 miles (600 kilometers) to the east, near Ames, Iowa. We knew that quaking and bigtooth aspen hybridize naturally within their range-- renowned forest ecologist Burton \"Burt\" Barnes's seminal work documented many such stands in Michigan (1961)--but the thought of this hybrid having occurred naturally so far outside the distribution of one of its putative parental species struck me, at least, as somewhat scandalous. Aspen pollen is wind-distributed and can travel long distances, but there is no evidence of successful pollination events occurring when around 375 miles separate male and female plants. As such, I expected that the reputation of the Smith Falls aspens as hybrids was nothing more than understandable wishful thinking. After all, bark and leaf traits can be plastic, and a more parsimonious explanation of the unusual appearance of the Niobrara trees was that they were simply an unusual, isolated stand of quaking aspen. So, an important first step in our research would be to compare the genetics and morphology of the Smith Falls aspens to that of known quaking and bigtooth trees. Our research group collected leaf samples, from which we could extract DNA, from all ten stands of Smith Falls aspens. We also dug up rhizomes to produce cloned suckers and planted these suckers in a common garden at a research station in Minnesota. There, we could perform experiments on them without harming the precious Niobrara trees. For the sake of comparison, we also drove all over the Midwest, collecting leaf and rhizome samples from quaking aspens in the Black Hills of South Dakota and the Sandhills of Nebraska, from bigtooth and quaking aspens along their western range edge in Minnesota and Iowa, and from both species within the interior of their distributions in Minnesota and Wisconsin. Our assessment of the genetics and physiology of the aspens took place over two years at the University of Minnesota campus. We used microsatellite genotyping--the same technology that allows for DNA fingerprinting in humans--to understand which trees we had sampled were distinct individuals and which were clones. This work, as well as sequencing of parts of our sampled trees' chloroplast genomes, was possible thanks to the full genome for black cottonwood (Populus trichocarpa), which was produced by an international team of dozens of biologists led by Oak Ridge National Laboratory's Gerald Tuskan in 2006--the first whole genome project carried out for a tree species. In our physiological experiments, we pushed stems and leaves from our common garden trees to the brink. We measured them exhaustively, and we then dried them out and froze them to mimic climate change-induced drought and post-budbreak freezes. We also tracked their phenology--the timing of their leafing out and loss of leaves in our common garden. An Ice Age relic The results of our genetics work (Deacon et al., 2017) left me picking my jaw up from the lab bench. Our first finding struck an ominous tone: the Smith Falls aspens are shockingly ARNOLD ARBORETUM, NATIONAL PARK SERVICE, NEBRASKA GAME AND PARKS COMMISSION, AND GIS COMMUNITY 8 Arnoldia 76\/3 ? February 2019 Genetic analysis of the ten aspen groves at Smith Falls (marked in red) revealed that only three genetic individuals occurred at the site. In some cases, these clones spanned opposite sides of the large canyon (and namesake waterfall) that bisects the park, suggesting aspens were once more widespread at the site. undiverse. We are confident that, across all ten stands at the site, any given tree belongs to one of three genotypes. This means that three original seedlings produced through sex have given rise, through rhizome suckering, to all of the extant aspens in the area. We found genetically identical individuals growing on opposite sides of ravines and in stands separated by hundreds of meters. It appears, then, that these particular trees rely almost exclusively on asexual suckering for reproduction. More shocking still, at the nuclear level, the Smith Falls aspens shared genetic information with both quaking aspen and bigtooth aspen, confirming that they are a hybrid between these parents--a hybrid christened, appropriately enough, Populus ? smithii. Evidence from chloroplast DNA suggested that bigtooth aspen, the species now not found until the middle of Iowa, was probably the maternal parent, with pollen coming from quaking aspen. Furthermore, patterns of genetic mixing we observed offer some support for the classification of these trees as F1 hybrids, meaning they are the first-generation offspring between two parents of different species, like mules. Our study of leaves collected from common garden trees supported our finding that the Smith Falls aspens were in fact hybrids of quaking and bigtooth parents (Deacon et al., 2017). Use of a dichotomous key to distinguish between these species will often require inspection of the pubescence and margins of the leaves of the specimen in question. Quaking aspens tend to be glabrous with many small teeth on their leaf margins. Bigtooth aspens tend to be pubescent with fewer, larger teeth. Barnes's work on Populus ? smithii tells the same story. Though we documented many subtle differences between leaves of the two species in our systematic analysis, our findings contributed to the current consensus: pubescence and tooth number are the best way to tell them apart. And Smith Falls Aspen 9 Specter of climate change Our grant from the National Parks Service enabled us to go beyond determining the genetic identity of these trees. We also used our common garden to study their disturbing demographic decline. State managers and conservationists had noted that existing aspens looked stressed and that new trees either were not sprouting or were quickly consumed by deer before outgrowing their reach. We wanted to understand the vulnerability of the Smith Falls aspens to two forms of physiological stress likely to be concomitant with climate change. The first is straightforward: climate change in the region is likely to lead to more arid conditions, imposing drought stress on the vulnerable, mesic species of the Niobrara River Valley. The second is less so: because the forces that cause this warming do not necessarily prevent late-winter cold snaps, even if spring temperatures arrive earlier, plants can leaf out in response to an early spring, then get hit with a freeze after budbreak. Such post-budbreak freezPHOTOS BY JAKE J. GROSSMAN leaves from P. ? smithii trees grown in Minnesota from Nebraska-collected rhizomes were perfectly intermediate between their putative parents in these two traits. Taken together, these findings suggest the rather shocking story that, at some point when both bigtooth and quaking aspen were locally abundant in the Niobrara River Valley-- probably between three and six thousand years ago--the two species hybridized. Both parents went locally extinct (and bigtooth vanished from the entire region), but their hybrid remained, reproducing vegetatively through rhizomes rather than through flowers and seeds. Despite considerable environmental change-- the climate in this region was warming and drying long before our present human-induced bout of climate change--these aspens hung on, perhaps shrinking in their distribution, but not disappearing from one small stretch of the Niobrara River Valley. As such, these aspens are a true relic of a past climate and a unique genetic treasure of the region. Each of the three aspen genotypes at Smith Falls is a first-generation hybrid, suggesting that the aspen trees found today at the site have descended through many generations of vegetative suckering, rather than seed reproduction, from the original hybrids. Vegetative suckering in aspens occurs via underground rhizomes, which the author showcases above. \"BUFFALO\" BRUCE MCINTOSH AND WESTERN NEBRASKA RESOURCES COUNCIL 10 Arnoldia 76\/3 ? February 2019 Large eastern red cedars (Juniperus viginiana) have encroached on the aspen groves at Smith Falls. This photo was taken before an intensive volunteer effort removed a large number of cedars, some shown in the midground of this forest. Ponderosa pine (Pinus ponderosa) rises in the background. ing can range from damaging to catastrophic, potentially killing vulnerable tissues and leading to whole-plant death (Anderegg et al., 2015). Our exploration of the aspens' vulnerability to climate change, currently in review, resulted in some bad and some good news. Unsurprisingly, given aspens' low level of drought tolerance, we found quaking, bigtooth, and hybrid aspens to be vulnerable to drought-induced cavitation, the formation of air bubbles in stem xylem. These bubbles, in quantity, disrupt waterflow in trees' vascular systems, like holes in a straw. They can, ultimately, lead to total hydraulic failure and tree death. Quaking aspen was slightly more vulnerable to this type of drought damage than bigtooth aspen, and their hybrid was intermediate. More generally, the hybrid aspens shared some drought-tolerance traits with quaking aspen, others with bigtooth aspen, and, in other cases, was intermediate between the two. But all three taxa showed a limited capacity to resist the challenges likely to occur in a warming and drying climate. The story surrounding post-budbreak freezing was simpler and rosier. We froze growing stems and leaves at temperatures equivalent to and lower than those that aspens in Nebraska are likely to experience during March and April storms. They were not substantially injured by this, suggesting that drought threatens aspens in the Niobrara River Valley more than latewinter cold snaps. Interestingly, in our measurements of spring phenology--the transition from dormancy to budbreak--we also found that the Smith Falls aspens were intermediate between quaking and bigooth aspen. Our findings echoed previous work showing that quaking aspens break bud about a week faster than bigtooths; fittingly, we observed that their hybrid offspring tended to leaf out in between the two parental species. Yet, compared to other regional conspecifics, all three groups of aspens generally leaf out around the same time. An aspen in a juniper's world Though our determination that the Smith Falls aspens are in fact hybrids has proven fascinating and satisfying, thornier questions remain about their future. Locally, the Smith Falls aspens are being outcompeted by neighboring trees, especially eastern red cedars (Juniperus virginiana). Quaking and bigtooth aspens are tolerant of low-intensity fires but intolerant of shade. Fire exclusion has, therefore, been catastrophic for \"BUFFALO\" BRUCE MCINTOSH AND WESTERN NEBRASKA RESOURCES COUNCIL Smith Falls Aspen 11 \"Buffalo\" Bruce McIntosh of the Nebraska Wildlife Federation leads a juniper removal initiative at Smith Falls. aspens, whether at Smith Falls or more broadly across the West. Because cedars are fire intolerant but drought tolerant, they have capitalized on our modern tendency to suppress and prevent fires. Locally, managers have fought this cedar encroachment by instigating prescribed burns, clearing cedars, and creating barriers to deer browsing on aspens. As a result, suckering is on the rise in some stands, producing a new cohort of healthy aspens (Robertson et al., 2018). Yet cedar removal and prescribed burns are expensive and will need to be repeated periodically to keep aspens abundant at Smith Falls. Globally, climate change is also likely to reshape the distribution of aspens across North America. Nebraska is expected to experience climate warming in the decades to come, in tune with the current global commitment to three to four degrees of warming. And though rainfall in the region will likely remain stable in absolute quantity, rain and snow will fall more sporadically, producing longer and more frequent periods of drought. None of this is good news for aspens in the region. Given historical changes in the distribution of quaking and bigtooth aspen, observations colleagues and I have made suggest the Smith Falls aspens will encounter a greater risk of climate-induced extirpation in the Great Plains than they have faced since the end of the last Ice Age. We might ask, then, what the future holds for the Smith Falls aspens and for other glacial relics in the region. The answer to this question depends on management. Stands of quaking aspen, paper birch (Betula papyrifera), and other drought-sensitive trees can probably be protected through active steps to shield them from direct climate stress and competition from more drought-adapted neighbors. In addition to prescribed burns, possible management practices include removal of competitors, use of exclosures to reduce grazing, targeted planting, and conservation of local groundwater. For species such as the aspens, which can be more easily propagated through rhizome cuttings than from seed, collection of rhizomes and propagation of suckers represents one pathway toward conservation of unusual, threatened germplasm. At present, private individuals and institutions may be able to assist with the migration of the Smith Falls aspens by purchasing commercially available nursery stock. Faller Landscape, in York, Nebraska, presently sells clonal trees produced through suckering from \"BUFFALO\" BRUCE MCINTOSH AND WESTERN NEBRASKA RESOURCES COUNCIL 12 Arnoldia 76\/3 ? February 2019 their flowers are fertile. Genetic evidence certainly suggests that no new trees have been born from seeds at the site for a very long time. And the Smith Falls aspens are beset by environmental challenges ranging from the hyperlocal to the global. Unfortunately, the Smith Falls aspens may simply have survived by demographic good fortune, constituting an evolutionary dead end, rather than a way forward. Yet this is a rather pessimistic view, and, having grown rather fond of the aspens at Smith Falls during my time working with them, I think their conservaThe fate of the Smith Falls aspens under climate change will likely parallel the fate tion is justified. As unusual of other drought-intolerant populations in the region, including this grove of quaking hybrids that have persisted aspen (Populus tremuloides), south of Smith Falls. despite millennia of climate rhizomes collected at the site (marketed under change--whether by chance or due to some the cultivar name `Ice Age'). 2 More broadly, particular adaptation we did not measure--they any efforts to stabilize the global climate, if our represent a potential genetic resource. As such, society is willing and able to undertake them, they may be candidates for assisted migration: will also benefit Midwestern aspens, among transplantation from their current, imperiled many other species. location to one that will be more appropriate in the coming centuries of a climate determined Hopeful monsters or dead ends? by human-induced changes. Propagation of Like other natural hybrids, the Smith Falls these trees in cooler and wetter climates well aspens have been heralded, at least locally, as within the current ranges of their parent species uniquely adapted to their surroundings. It is might allow them to flourish while also buytempting to assert that these trees, by virtue ing time for intentional propagation of secondof their longevity, may illustrate one strategy generation hybrids through breeding with other for persistence in a warming, drying climate. Popolus ? smithii or backcrossing. Yet findings from our research do not support Regardless, we would be wise to remember this narrative. Indeed, these hybrids are unique, that the story of evolution in response to a and worthy of study insofar as they constitute warming climate has occurred many times in an evolutionarily unusual relic from a past the history of life, and it often produces messy climate. But as noted above, these trees repstories like that of the Smith Falls aspens. What appear to us to be non-adapted (or even resent essentially three genetic individuals maladapted) trees may hold the key to survivthat have probably been cloning themselves ing and thriving in a future climate. After all, for thousands of years. Though we have very the common ancestor of modern aspens and recently received second-hand confirmation willows had likely evolved during the earth's that the trees do flower in some years, we do last period of extreme warming, some fifty-five not know the sex of each clone, or whether Smith Falls Aspen 13 million years ago, when the tropics extended up to the North Pole (Manchester et al., 2006). Though the fate of quaking and bigtooth aspens and their hybrids is uncertain, the aspen lineage is likely to survive contemporary climate change. And so, whether hybrid aspens are best thought of as hopeful monsters or evolutionary dead ends is ultimately unknowable. But as an unlikely yet arguably successful hybrid myself, I'm inclined to give them the benefit of the doubt. Acknowledgements The original research reported in this article was conducted collaboratively at the University of Minnesota--Twin Cities with Jeannine Cavender-Bares, Nick Deacon, Anna Schweiger, and Isabella Armour. We were funded by National Park Service grant #191779 to Jeannine Cavender-Bares, Mark Dixon, and Molly Nepokroeff. Jeffrey Carstens, a horticulturist and curator of woody and herbaceous plants at the USDA's North Central Regional Plant Introduction Station in Ames, Iowa, played an absolutely essential role in guiding our collections of Midwestern aspens, especially along the species' range edges in Iowa. \"Buffalo\" Bruce McIntosh of the Nebraska Wildlife Federation and Joseph Zeleznik at North Dakota State University also provided essential help with collections in Nebraska and northern Minnesota, respectively. Three photographs in this article came from the video \"Fighting for Survival: The Ancient Aspen of the Niobrara Valley,\" produced by \"Buffalo\" Bruce McIntosh, Rod Jensen, and The Great Plains Motion Picture Company. Notes 1It should be noted that the chimaeras resulting from the almost unbelievable process of interspecies grafting (e.g. apples, pears, grapes, roses, citrus, mangos, stone fruit, and others, not to mention the introduction of certain porcine organs into human bodies) certainly give more integrated hybrids a run for their money in the realm of public interest. 2Aspen enthusiasts can also purchase from them a very robust and aesthetically pleasing quaking aspen genotype, `NE Arb', which was collected from a nowextinct stand elsewhere in Nebraska. References Anderegg, W.R.L., A. Flint, C.Y. Huang, L. Flint, J.A. Berry, F.W. Davis, J.S. Sperry, and C.B. Field. 2015. Tree mortality predicted from droughtinduced vascular damage. Nature Geoscience, 8: 367?371. Barnes, B.V. 1961. Hybrid aspens in the Lower Peninsula of Michigan. Rhodora, 63: 311? 324. Bessey, C.E. 1887. A meeting-place for two floras. Bulletin of the Torrey Botanical Club, 14: 189?191. Burger, W.C. 1975. The species concept in Quercus. Taxon, 24:45?50. Deacon, N.J., J.J. Grossman, A.K. Schweiger, I. Armour, and J. Cavender-Bares. 2017. Genetic, morphological, and spectral characterization of relictual Niobrara River hybrid aspens. American Journal of Botany, 104: 1878?1890. Goldschmidt, R. 1940. The material basis of evolution. New Haven, CT: Yale University Press. Hamzeh, M., and S. Dayanandan. 2004. Phylogeny of Populus (Salicaceae) based on nucleotide sequences of chloroplast trnT-trnF region and nuclear rDNA. American Journal of Botany, 91: 1398?1408. Kaul, R.B., G.E. Kantak, and S.P. Churchill. 1988. The Niobrara River Valley, a postglacial migration corridor and refugium of forest plants and animals in the grasslands of central North America. The Botanical Review, 54: 44?81. Manchester, S.R., W.S. Judd, and B. Handley. 2006. Foliage and fruits of early poplars (Salicaceae: Populus) from the Eocene of Utah, Colorado, and Wyoming. International Journal of Plant Science, 167: 897?908. Robertson, J.M., A.R. Cahlander-Mooers, and M.D. Dixon. 2018. Effects of management treatments on regeneration of a geographically disjunct, relictual hybrid aspen (Populus ? smithii) population in the central Great Plains, USA. Environmental Management, 62: 906?914. Tuskan, G.A, S. Difazio, S. Jansson, J. Bohlmann, I. Grigoriev, U. Hellsten,... D. Rokhsar. 2006. The genome of black cottonwood, Populus trichocarpa (Torr. & Gray). Science, 313: 1596?604. Wright, H.E., J.C. Almendinger, and J. Gr?ger. 1985. Pollen diagram from the Nebraska Sandhills and the age of the dunes. Quaternary Research, 24: 115?120. Wu, G.A., J. Terol, V. Ibanez, A. L?pez-Garc?a, E. P?rez-Rom?n, C. Borred?,... M. Talon. 2018. Genomics of the origin and evolution of Citrus. Nature, 554: 311?316. The maps in this article were created using ESRI, USGS, USFS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, Intermap and the GIS user community. Jake J. Grossman is a Putnam Fellow at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Roaming through Ranges: The Evolution of Tree Species Distribution Maps in the United States","article_sequence":2,"start_page":14,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25656","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060bb26.jpg","volume":76,"issue_number":3,"year":2019,"series":null,"season":null,"authors":"Ellenwood, James","article_content":"ELLENWOOD, J. 2019. ROAMING THROUGH RANGES: THE EVOLUTION OF TREE SPECIES DISTRIBUTION MAPS IN THE UNITED STATES. ARNOLDIA, 76(3): 14?27 Roaming through Ranges: The Evolution of Tree Species Distribution Maps in the United States James Ellenwood A s a teenager, in the 1970s, I traveled with my siblings across the United States and Canada on a few multi-week road trips, camping and visiting National Parks and Forests. Along the way, I became fascinated by how different trees grow in different places-- the tall, narrow-crowned Engelmann spruce (Picea engelmannii) found in the high elevations of the northern Rocky Mountains (which were still snow packed in the middle of June) and the incense cedar (Calocedrus decurrans) showing off its brilliant red bark on the western slope of the Sierras. The local tree species, I realized, were as distinct as the terrain-- patterns that emerged over millions of acres and thousands of miles. The quest for adventure and the desire to see the natural world drove my ambitions, facilitated by recently completed interstate highways and a personal automobile. Ecological patterns that were once relatively difficult to see now emerged, much like the transition from still frames to movie frames, via easy roadside stops to get into the wilds. Around this exact same time, these fundamental biological patterns were being synthesized into a comprehensive set of North American tree distribution maps by a botanist at the United States Forest Service named Elbert Little Jr. These maps were published in a six-volume set, collectively titled the Atlas of United States Trees, which covered around 720 species. Each map included shapes that represented the maximum extent of the distribution for a single species. Little and his collaborators developed these by drawing an outline around locations obtained from numerous published and unpublished sources, as well as personal knowledge. For decades, Little's maps have served as the definitive source for this distribution information. I referenced them as a student at the State University of New York's College of Environmental Sciences and Forestry in the early 1980s. And digital reproductions of Little's maps are now included on the Wikipedia pages for most--if not all--of the tree species that he documented. In 2007, however, I began working with colleagues at the United States Forest Service to project the potential for forest mortality due to insects and diseases fifteen years into the future. This involved building thousands of models based upon remotely sensed imagery, environmental variables, and observations from field plots. After many trials and iterations, we recognized a need to bound the modeled maps by known tree distributions, but we knew that Little's maps couldn't be used for this type of analysis. For instance, if we were attempting to understand the threat of emerald ash borer (Agrilus planipennis) on populations of white ash (Fraxinus americana)--a widespread forest species in eastern North America--Little's map would suggest that white ash was equally prevalent in New York and Illinois, given that both states fall entirely within the range. Yet New York is widely forested, while Illinois is extensively cultivated. As a result, white ash populations would be much more fragmented in the western portion of its range, and Little's maps did not reveal this trend. Clearly new information was needed. Original Maps The history of mapping North American tree distributions is an ongoing narrative of collaboration on a national scale, but the evolution of these maps also reveals how cultural values (and even politics) factor into seemingly straightahead descriptive botany. My team began working on new distribution maps because of environmental concerns about the globalization of insects and diseases, an issue that wasn't remotely part of the scientific consciousness in 1880, when Charles Sprague Sargent, the founding director of the Arnold Arboretum, was leading an initiative to describe and inventory UNITED STATES FOREST SERVICE, FOREST HEALTH TECHNOLOGY ENTERPRISE TEAM Species Distribution Maps 15 This map for white ash (Fraxinus americana), published in the National Individual Tree Species Atlas (2015), compares a classic distribution prepared by Elbert Little Jr. (light purple) with a new modelled distribution (dark purple). forested landscapes for the 1880 United States Census. Rather, at that point in time, the primary driver for this kind of distribution mapping was related to economic interests. Sargent compiled information for about 412 tree species, which appeared in a voluminous final report, published in 1884. Some of this information came through personal field observations, but Sargent also enlisted the support of a team of botanists and natural history enthusiasts who traveled through specific regions, reporting on the composition of the forests. Most of the resulting distribution information was text-based, but the report also included several types of maps: One set showed the extent of forested landscapes for individual states. Another set, included as a large-format portfolio, showed the distribution of genera, 16 Arnoldia 76\/3 ? February 2019 more detailed representation, and the maps also indicated \"regions from which Mercantile Pine has been removed.\" Farther west, individual species were not tracked on the state maps, which instead showed the density of forests. Likewise, the supplementary map for all pines indicated only the density (essentially a heatmap) rather than showing the ranges for individual species. As such, if someone wanted to determine the scattered range for something like the whitebark pine (P. albicaulis), which favors subalpine regions in the western mountains, the text-based descriptions would have been the best resource. Of course, individual species maps would have required many more ARNOLD ARBORETUM ARCHIVES like pines (Pinus) or oaks (Quercus), on a continental scale, with monochromatic shading used to indicate the number of species in each location. All the maps were prepared by Andrew Robeson, Sargent's brother-in-law. It is particularly interesting to look at how the census report renders a single economically important species, like the long-leaf pine (Pinus palustris). Sargent called the species \"a tree of first economic value\" and described a distribution that spanned from southeastern Virginia, through the Gulf states, and into eastern Texas, \"rarely extending beyond 150 miles from the coast.\" On maps of the corresponding states, long-leaf pine was singled out for Charles Sprague Sargent's 1884 census report included forest maps for individual states. For South Carolina, the distribution of two species of long-leaf pine (both now recognized as Pinus palustris) are shown in green. Commercially harvested forests (shown in burnt orange) emanate from the coast and along transportation corridors. ARNOLD ARBORETUM ARCHIVES Species Distribution Maps 17 The census publication included sixteen large-format maps, including one showing the density of pine species. The darkest shading, shown only in southern California, indicates that the census recognized seven pine species in that area. field observations than were currently available. A large region in the center of the Idaho census map was still labelled \"unexplored,\" which now offers an evocative reminder of this information scarcity. While the census project was nearing completion, Sargent simultaneously chaired an initiative to preserve woodlands in New York's Adirondack Mountains from deforestation, which culminated in the establishment of Adirondack Park--the country's first state forest preserve--in 1885. It is clear that Sargent was aware of the utilitarian value of the cen- sus project, given that more than two hundred pages of the final report were devoted to the material properties of wood derived from each species, yet mapping the forests also revealed the finite dimensions of a resource that had once seemed limitless, potentially setting the stage for subsequent conservation and preservation movements. Developing Detail While the census report provided a useful synthesis of information known at the time, by 1898, George Bishop Sudworth, a dendrologist 18 Arnoldia 76\/3 ? February 2019 for the United States Department of Agriculture's Division of Forestry, articulated the need for a more comprehensive and up-to-date treatment of distribution information. \"The army of professional and amateur botanists engaged in botanical research are yearly bringing to light new facts, which are constantly enlarging our understanding of the geographical distribution of trees and other plants,\" Sudworth wrote in the first edition of his Check List of the Forest Trees of the United States: Their Names and Ranges, which was published that year. The checklist included short descriptions of the ranges for five hundred species, but he knew that much more collaborative work was still needed. The 1898 checklist ultimately presaged Sudworth's lifelong effort to develop better species distribution maps. Significantly, in 1913, Sudworth published an atlas for North American pine species, which was intended to be the first volume of a series covering all native trees. These maps represented a significant step towards the familiar appearance of distribution maps today, with each species rendered on a single map showing the entire range. Sudworth even showed the distribution of relatively sparse species like the whitebark pine, with green marking that followed the narrow elevational bands along mountain ranges. Given the scale, a reader would have difficulty in seeing the range for whitebark pine without the use of visual aids such as a magnifying glass, but the effort suggests the amount of detailed field observation that went into the project. Sudworth attributed the success of his maps to work being conducted by the new United States Forest Service, established in 1905, which provided invaluable \"unpublished field notes, unrecorded observations, and reports of Forest Service officials engaged in the exploration, surveying, and administration of the 163 National Forests now established.\" Sudworth's assistants compiled information from these sources, along with state floras and other resources, on cards for each species, and these annotations were plotted on a map of North America. Sudworth knew that the simplicity of maps far exceeded the usefulness of even the most detailed text, yet his volume on pines was the only portion of the atlas ever published. The explanation for Sudworth's discontinuation of the atlas project might be intuited from the publication of subsequent bulletins that included maps of select tree species (including additional conifers) in the Rocky Mountains. Given that much of Sudworth's field information arrived from the National Forests, which were almost entirely located in western states (at the time), it makes sense that the maps would ultimately share the same regional emphasis, with special attention given to species that were of economic importance. In 1927, the year of Sudworth's death, he published a revised checklist of North American tree species, in which he briefly described the range for every known species at the time (adding more than three hundred taxa to his original checklist). Sudworth knew the information would prove useful \"not only among foresters, woodsmen, and wood users, but also in forest schools and other educational institutions.\" At the time, he and an assistant were once again working on maps--with many complete but unpublished--but the effort was suspended as priorities shifted (perhaps related to the Great Depression). Developing Breadth Edward Norfolk Munns, the chief of the Forest Service's Division of Forest Influences, eventually returned to the work that Sudworth and his assistants had started. In 1938, he published an atlas covering 170 of the most important tree species, which he noted was \"based very largely\" on Sudworth's research, with many updated observations compiled by junior forester William W. Mitchell. Although the species representation was far greater than anything published in Sudworth's lifetime, critics suggested that the maps should have been shared with field botanists and foresters for additional corroboration, because to many working on the ground, errors were evident. Even so, the atlas was reprinted by popular demand. Beyond utility for botanists, foresters, and \"the manufacturer in search of raw materials,\" Munns also described new ecological and engineering implications for the maps, no doubt based on his early field experience studying the impact of wildfires on California watersheds. BIODIVERSITY HERITAGE LIBRARY, CONTRIBUTED BY USDA NATIONAL AGRICULTURAL LIBRARY Species Distribution Maps 19 George Bishop Sudworth's 1913 atlas included distribution maps for thirty-six species of pines. This map shows the scattered high-altitude distribution of whitebark pine (Pinus albicaulis) BIODIVERSITY HERITAGE LIBRARY, CONTRIBUTED BY USDA NATIONAL AGRICULTURAL LIBRARY 20 Arnoldia 76\/3 ? February 2019 This detailed rendering of long-leaf pine (Pinus palustris) appeared in the 1938 atlas prepared by Edward Norfolk Munns. \"[Forest distribution] is an essential element in erosion and flood control operations, and in land-use planning,\" he wrote in the introduction. \"Indeed, present trends toward better planning and integration of land use are directing increased attention to forest cover, the species represented in it, and the possibilities of enlarging the contribution of forest land to community welfare.\" This, of course, may have been a suggestion that worked better on paper than in practice, but the statement represented an important expansion of what distribution maps could enable. Ultimately, in 1942, a forest ecologist name Elbert Little Jr. was appointed as dendrologist for the Forest Service. Like both Sudworth and Munns (and myself), Little had spent a considerable amount of time at Forest Service field stations in the West, before taking his appointment at the national office in Washington. Little's continuation of the mapping project began with small generalized ranges for 165 forest species of economic interest, published in 1949. He later explained that the small size of the maps was due to practical and logistical reasons, given that it is easier to approximate an accurate range at a smaller scale. \"Botanists, foresters, and other authors bold enough to summarize plant distribution records graphically may expect criticism instead of reward for their efforts,\" he wrote in a follow-up article in Rhodora. \"It is far easier to detect a minor flaw along a boundary line than to prepare a better map.\" I learned this lesson very quickly in my own endeavors, to say the least. Over the next two decades, Little and his assistants worked rigorously to expand the distribution data, sifting through more than three hundred sources, including unpublished card files in state herbaria and doctoral dissertations. Like Sudworth's maps, the reference points were then plotted onto a map of the United States or Species Distribution Maps 21 transparencies, the originals could be used with two of the other volumes. If Munns, therefore, alluded to the possibility of using the distribution maps for something beyond a guide to natural resources, Little's transparencies indicated a genuine commitment to expanding the types of questions that could be raised with the maps. \"They provide the basis for correlation studies of distribution of a species and the environment,\" Little wrote of the overlays (in the fourth volume). Moreover, as someone who began his career as a forest ecologist, Little saw a greatly expanding set of research questions that would benefit from the maps, including \"such studies as classification, evolution, paleobotany, and genetics, and for the distribution of associated animals and plants, especially insects and parasitic fungi.\" Although the transparencies may seem simple compared to modern approaches, the effort to enable comparisons between tree BIODIVERSITY HERITAGE LIBRARY, CONTRIBUTED BY USDA NATIONAL AGRICULTURAL LIBRARY North America. The first volume appeared in 1971, and the next five volumes appeared over the next ten years, ultimately covering more than seven hundred species of trees and major shrubs. When Little published on the distribution of trees in Alaska, he retained the individual reference points within the distribution outlines, but otherwise, the maps generally followed the classic form, with a simple outline drawn around all contiguous populations. This work still represents the standard reference for tree species ranges in North America today. The first volume of the atlas also included nine semi-transparent overlays, nested within a cover pocket, which could be superimposed over the range maps. The overlays included features like \"precipitation and rainfall,\" \"plant hardiness zones,\" and \"maximum extent of glaciation in the Wisconsin Glacial Stage.\" While the subsequent volumes did not include these For many water-loving species, like the black willow (Salix nigra), Munns rendered the distribution according to rivers and streams. 22 Arnoldia 76\/3 ? February 2019 species and environmental conditions represents a milestone development. Once published, Little worked vigorously in trying to maintain an authoritative record for each individual species, even after retirement. A colleague of his related a story of visiting an area in Wisconsin in the 1980s where there was a rumor for the occurrence of a species that wasn't depicted in the atlas. Little brought his field maps, confirmed the sighting, and promptly penciled in the location. Eventually, the field maps were brought back to the office for inclusion in the authoritative maps. Little stressed the importance of continual maintenance. His coauthor on the second volume, Leslie Viereck, continued to maintain records for Alaska and produced a second edition, which included range adjustments, some species reclassifications, and an expansion to include significant shrub species. It was unfortunate that Viereck passed away in August of 2008, about a year before we started on the finer scaled species distributions. Modelling the Present After Little retired from the Forest Service in 1975, the position of dendrologist was unfortunately abolished. In the ensuing decades, ecosystem classifications became the mapping priority. Communities of trees were identified as the dominant factors necessary for analyzing impacts of forest management, and distribution maps for individual tree species were no longer emphasized. With shifting priorities, staff changes, and a relocation of the National Headquarters, the whereabouts of Little's authoritative maps was lost through the ages. Although I have encountered people whose tenures overlapped with Little's, it is likely that the data have been forgotten, left in an attic, or moved to a storage facility at the Department of Agriculture's Greenbelt center or to a National Archives and Records Administration facility. If these data are recovered in the future, it would be important to appropriately curate them for further refinement and research. In some disciplines, however, the need for better species maps became critical. Given practical considerations, Little was required to take inherent liberties when connecting a distribution outline around scattered dots, and the resulting shapes also failed to convey the density of forests. Moreover, the maps no longer represented the best information about the current distributions. Limitations like this are inherent to the mapmaking process. Even Sudworth, writing in 1913, noted the forests were changing faster than the maps could show. \"Extensive and continued lumbering operations with attending forest fires have so changed, and in some cases exterminated, parts of the original stand of most of our pines,\" he wrote. \"These maps, therefore, indicate only the general occurrence of species with the prescribed areal limits, and have no reference to the density or continuity of growth.\" Munns made the same point twenty-five years later, and Little echoed these concerns. These limitations ultimately fueled my work with the Forest Service's Forest Health Technology Enterprise Team, where we needed to develop models to predict the risks associated with forest pests. We obtained current observations of 346 species that occur at Forest Inventory and Analysis field plots, as well as from other permanent plots managed by the National Forest System and the Bureau of Land Management. This amounted to more than 330,000 plots (and more than 1.2 million subplots). We linked this information to predictive layers pertaining to environmental variables like climate, terrain, soils, and satellite imagery. This dataset was then used to model individual species presence, as well as stand density, which was necessary for pest risk mapping. In the end, we successfully modelled the distribution of 264 trees sampled on these Forest Service plots, and as a by-product of that work, we published the National Individual Tree Species Atlas in 2015. Of course, this printed document will ultimately become a historical artifact, much like Little's volumes, given that the forests in the country will continue to change and transform in response to disease and insect pressure, climate change, habitat destruction, disturbance recovery, and any number of other threats (or boons), many still unforeseen. The difference between the distributions shown on Little's maps and our models is often quite noteworthy, so our published atlas also ARNOLD ARBORETUM ARCHIVES Species Distribution Maps 23 The first volume of Elbert Little Jr.'s Atlas of United States Trees (1971) included nine overlays that allowed for environmental analysis. In this case, a topographical overlay has been superimposed over the range of whitebark pine (Pinus albicaulis). Also note that the atlas included county borders. UNITED STATES FOREST SERVICE, FOREST HEALTH TECHNOLOGY ENTERPRISE TEAM 24 Arnoldia 76\/3 ? February 2019 This map of long-leaf pine (Pinus palustris), published in the National Individual Tree Species Atlas, shows a smaller modeled distribution (dark purple) compared to the distribution outlines prepared by Elbert Little Jr. (light purple). includes Little's outlines for comparison. It was my hope that this format would draw awareness to the ongoing need for studying these ranges--work that might require the oversight of a twenty-first-century Little (an authoritative steward of tree species distribution data)-- however, even now, no specific authority is responsible for maintaining comprehensive distribution records. The United States Department of Agriculture's Natural Resources Conservation Service maintains the PLANTS Database, which has some degree of authority, although species distributions are only tracked at the county level, at best, which is adequate for general applications but not for applications in need of a finer scale. Species Distribution Maps 25 For many species, like the long-leaf pine, comparison between the models and Little's maps suggests range contraction. The same is true for the whitebark pine. Both species are recognized as endangered, according to the International Union for Conservation of Nature, although for different reasons. The long-leaf pine was eventually disfavored by the forest industry due to its lengthy sapling (\"grass\") stage, and as a result, commercially managed forests were preferentially replanted with loblolly and slash pines (Pinus taeda and P. elliottii, respectively). The species is still threatened with continued habitat loss, although substantial restoration efforts are underway. Whitebark pine is currently most threatened by recent outbreaks of mountain pine beetle (Dendroctonus ponderosae), a native insect that has caused widespread mortality among western pine forests. In both cases, the models are critically important for monitoring current populations, as well as for projecting the future of these populations. Other species, like the Osage orange (Maclura pomifera), however, show a dramatic range expansion. The range shown on Little's map is an upright column running through eastern Texas, barely extending into southern Oklahoma and Arkansas. Little wanted to show the original range for the species, before it had been widely planted as a living fence between agricultural fields in the Midwest. Because the species readily naturalized, our models, based on information about actual occurrence at field plots, shows a much wider range, with populations as far afield as western Pennsylvania. Our atlas also includes ranges for three nonnative species--tree of heaven (Ailanthus altissima), Chinese tallow tree (Triadica sebifera), and empress-tree (Paulownia tomentosa)--which have naturalized widely. Since these ranges weren't recorded in Little's atlas, this will provide invaluable baseline information for future management and research efforts. Changes to species classification can also result in significant changes to the distribution maps. The bristlecone pine, for instance, was separated into two species--Rocky Mountain bristlecone pine (Pinus aristata) and Great Basin bristlecone pine (P. longaeva)--which were easily separated based upon geographic data. Other species classification changes were not so easy. The Mexican pinyon pine (P. cembroides) was divided into two additional species, border pinyon (P. discolor) and papershell pinyon (P. remota), but the distributions were much more difficult to separate due the coincidence of the three species. As the inventory is maintained, newer modeling techniques can improve the distribution maps for only the species measured on an inventory plot. Other naturally occurring species--often those with more restricted ranges in the first place--will need different data sources and greater effort to be developed. Beyond Borders While developing our models, we had the privilege of working with colleagues in Mexico to develop pest risk maps for Douglas fir (Pseudotsuga menziesii) and several key pines. Though we limited our investigation to eight species that had coarse climate and soils data, the permanent inventory for Mexico is designed much like the United States, and the potential exists to develop a complete set of species distribution maps for Mexico. (It should be noted that while 387 species were encountered within inventory plots in the United States, the Mexico inventory counted over 3,000 tree species.) Meanwhile, the resolution of our information does not carry into Canada, given limited access to the same amount of field data. Canada produced distribution maps for approximately ninety-three species, although they are of limited precision compared to the maps in the United States. At present, however, our own models don't extend north of the border either. In this sense, political relationships are often implicit in distribution maps, much as economic and ecological imperatives have manifest themselves throughout this ongoing history. Given the nature of remote sensing, however, there is increasing potential to combine forest inventories to map complete species distributions, regardless of political boundaries. The North American Forest Commission is currently developing a combined database for Mexico, Canada, and the United States, and the success of a shared system like this was recently 26 Arnoldia 76\/3 ? February 2019 consistent methods and metrics), the resulting atlas is testament to the achievements possible with international collaboration. Moreover, the Food and Agriculture Organization of the United Nations asks all countries to assemble a National Forest Inventory every five years, and although the distribution data are relatively UNITED STATES FOREST SERVICE, FOREST HEALTH TECHNOLOGY ENTERPRISE TEAM demonstrated in Europe. The European Union published the first systematic atlas of trees at the continental scale in 2016, which grew out of an effort to harmonize data within a continentwide forest information system, established in 2013. Although the authors stressed the need for even more data (collected using more This map of Osage orange (Maclura pomifera), published in the National Individual Tree Species Atlas, shows an expanded modeled distribution (dark purple) compared to Little's distribution outlines (light purple). Species Distribution Maps 27 coarse, efforts like this suggest the potential for a much more comprehensive set of tree species maps, especially in temperate regions where species diversity is less complex. Taken together, these aspirations suggest the longevity of Sudworth's observations in 1898. \"The geographical range of any of our trees must necessarily be an expression of the united efforts of all working botanists,\" he wrote, \"for the unaided diligence of one man's lifetime could never carry his search and study into all of nature's hiding places for even trees alone.\" Since the completion of the National Individual Tree Species Atlas, many changes have occurred that would enhance future modeling efforts. Modeled maps now have the potential to be dynamic and adaptive, but they still require the collaborative vision of botanists, foresters, and plant ecologists in the field, now and for generations to come. References: Beaudoin, A., Bernier, P.Y., Villemaire, P., Guindon, L., and Guo, X.J. 2017. Species composition, forest properties and land cover types across Canada's forests at 250m resolution for 2001 and 2011. Quebec: Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre. Ellenwood, J.R., Krist F.J., and Romero, S.A. 2015. National individual tree species atlas. Washington, DC: United States Forest Service, Forest Health Technology Enterprise Team. Little, E.L. 1949. Important forest trees of the United States. In Trees: The yearbook of agriculture. Washington, DC: United States Department of Agriculture. Little, E.L. 1951. Mapping ranges of the trees of the United States. Rhodora, 53: 195?203. Little, E.L. 1971. Atlas of United States trees: Conifers and important hardwoods (Vol. 1). Washington, DC: United States Department of Agriculture Forest Service. Little, E.L. 1976. Atlas of United States trees: Minor western hardwoods (Vol. 3). Washington, DC: United States Department of Agriculture Forest Service. Little, E.L. 1977. Atlas of United States trees: Minor eastern hardwoods (Vol. 4). Washington, DC: United States Department of Agriculture Forest Service. Little, E.L. 1978. Atlas of United States trees: Florida (Vol. 5). Washington, DC: United States Department of Agriculture Forest Service. Little, E.L. 1981. Atlas of United States trees: Supplement (Vol. 6). Washington, DC: United States Department of Agriculture Forest Service. Munns, E.N. 1938. The distribution of important forest trees of the United States. Washington, DC: United States Department of Agriculture Forest Service. De Rigo, D., Caudullo, G., Houston Durrant, T., and San-Miguel-Ayanz, J. 2016. The European atlas of forest tree Species: Modeling, data and information on forest tree species. In: San-Miguel-Ayanz, J., De Rigo, D., Caudullo, G., Houston Durrant, T., and Mauri, A. (Eds.), European atlas of forest tree species. Luxembourg: Publishing Office of the European Union. Sargent, C.S. 1884. Report on the forests of North America (exclusive of Mexico). Washington, DC: Department of the Interior Census Office. Sudworth, G.B. 1898. Check list of the forest trees of the United States: Their names and ranges. Bulletin of the United States Department of Agriculture, Division of Forestry, 17: 1?144. Sudworth, G.B. 1913. Forest atlas: Geographic distribution of North American trees. Part 1: Pines. Washington, DC: United States Department of Agriculture Forest Service. Sudworth, G.B. 1927. Check list of the forest trees of the United States: Their names and ranges. United States Department of Agriculture Miscellaneous Circular, 92: 1?297. Viereck, L.A., and Little, E.L. 1975. Atlas of United States trees: Alaska (Vol. 2). Washington, DC: United States Department of Agriculture Forest Service. Viereck, L.A., and Little, E.L. 2007. Alaska trees and shrubs (2nd ed.). Fairbanks, AK: University of Alaska Press. James Ellenwood is the National Program Lead for Monitoring, Remote Sensing, and Geospatial Analysis Research at the United States Forest Service. "},{"has_event_date":0,"type":"arnoldia","title":"Taiwan Dispatches","article_sequence":3,"start_page":28,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25658","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060816f.jpg","volume":76,"issue_number":3,"year":2019,"series":null,"season":null,"authors":"Wilson, E. H.","article_content":"WILSON, E.H. 2019. TAIWAN DISPATCHES. ARNOLDIA, 76(3): 28?33 Taiwan Dispatches Ernest Henry Wilson In March of 1919--one hundred years ago--Ernest Henry Wilson returned from his sixth and final plant collecting expedition to eastern Asia. The trip had begun in Yokohama, Japan, in January of 1917, and he traveled widely, tracing his way from Okinawa to Korea, even touching briefly into China. In early 1918, he sailed for Taiwan, where he was enamored with the subtropical conifers. According to his own tally, he collected more than seven thousand herbarium specimens, and he would return for seed in the fall. Taiwan, then known as Formosa, had been occupied by Japanese troops for more than two decades. Wilson's travels were conducted with a Japanese botanist named Ryozo Kanehira, and initial collecting locations were recommended by Bunzo Hayata, a botanist at the Imperial University of Tokyo. The following excerpts come from Wilson's handwritten letters to Charles Sprague Sargent, the director of the Arnold Arboretum. Italics have been added. JANUARY 17, 1918 | YOKOHAMA, JAPAN My arrangements are all completed for the trip to Formosa & I leave here in the morning. I plan to stay in Formosa for about ten weeks if the money in hand be enough to enable me to do so. JANUARY 25, 1918 | TAIPEH, FORMOSA Just a note to let you know that I have reached Formosa & that everything is favorable to a successful visit. I arrived here on the 22nd & leave tomorrow (26th) for Arisan where the giant trees are.... The officials, one & all, promise every assistance & there is no doubt but that they mean it. A Mr. Kanehira, who speaks English & is one of the heads of the forestry department, has been detailed to accompany me to Arisan & will probably go elsewhere with me also. He is a very nice fellow & I fancy will make a genial companion.... It is now eleven years since I begged some scraps of Taiwania [a monotypic member of the cypress family (Cupressaceae)] from Hayata & got promises, which were never fulfilled, of more material of Formosan conifers. I intend now to make up for lost time & our Herbarium shall possess its compliment of Formosan conifers ere I am through. FEBRUARY 16, 1918 | TAIPEH, FORMOSA I am back from the trip to Arisan & have but one regret which is that you too were not present to enjoy the forests & the giant trees. I had expected much but what I saw far exceeded my expectations: the forests are easily the finest & the trees the largest I have ever seen.... The country is very steep & savage & travelling over it is hard work. Thanks to a light railway & courtesies extended by the government things were made as easy for me as they possibly could be made. The weather on the whole was good though two consecutive days of rain & sleet & many foggy afternoons were a hinderance. I collected over twelve hundred specimens, representing about two hundred species, & took six and one-half-dozen photographs ... The Chamaecyparis formosensis [an endemic false cypress] is the largest tree being sometimes ALL IMAGES FROM ARNOLD ARBORETUM ARCHIVES Wilson's Letters 29 Wilson was awed by the enormous conifers near Mount Arisan. On February 1, 1918, he photographed Taiwania cryptomerioides (at right), towering beside Taiwan cypress (Chamaecyparis formosensis, at left). 30 Arnoldia 76\/3 ? February 2019 The botanist Ryozo Kanehira accompanied Wilson in Taiwan. On January 31, 1918, Wilson photographed Kanehira near Mount Arisan, standing beside the trunk of Taiwania cryptomerioides, which soared to a height of 150 feet (46 meters). Wilson's Letters 31 Clockwise from upper left: Lithocarpus amygdalifolius on February 1, 1918; Taiwan Douglas-fir (Pseudotsuga sinensis var. wilsoniana) on April 6; Taiwan cypress (Chamaecyparis formosensis) with a trunk diameter of 20 feet (6 meters), photographed on the return trip, October 31; and Calocedrus formosana on April 1. 32 Arnoldia 76\/3 ? February 2019 Wilson photographed pure stands of Taiwan fir (Abies kawakamii) on Mount Kiraishu, Nantou County, on March 6, 1918. nearly 200 ft. tall & 65 ft. in girth of trunk.... I was informed that the oldest tree which had been felled showed about two thousand seven hundred annual rings, & a larger one standing is estimated at three thousand years. The trunks are mostly hollow but the wood, which is reddish, fragrant, & has a beautiful satiny luster, is much esteemed by Japanese for interior work in houses. FEBRUARY 28, 1918 | TAIPEH, FORMOSA I returned from the trip to the south on the night of February 26th. The flora of the coastal region did not prove at all interesting, indeed, most of it had been destroyed to make way for sugar, rice, & other crops. However, I made a fair collection of plants & took a dozen photographs so we shall have a record of what the flora is like. MARCH 16, 1918 | TAIPEH, FORMOSA The trip to the central range of Formosa has proved a complete success. The weather was fine throughout & the journey fairly easy. From the railway we travelled for two days on push trolley & then climbed for three days, sleeping in police huts at night.... The peak we ascended is named Mt. Kiraishu, is 11,002 ft. high, well-forested on the upper-middle slopes. The climate is drier & the flora different from Wilson's Letters 33 that of the Arisan region. Abies kawakamii [an endemic fir] was the particular quest of the trip & we found it in great plenty above 9,500 ft. After collecting from the ground scales & spikes of disintegrated cones, I was fortunate enough to find four or five perfect cones & so complete the specimens. With this acquisition our herbarium possesses ample & complete material of every known species of Abies found in the Far East.... Altogether the trip yielded about two hundred species bringing the total to date collected in Formosa to about four hundred & seventy species. It was difficult country to photograph in but I secured two & one half dozen, which will give a fair idea of the vegetation. APRIL 11, 1918 | TAIPEH, FORMOSA I am writing this at the completion of the allotted task in Formosa. One objective I had in mind on visiting the island was to see if possible every conifer known to grow there. Dr. Hayata in Tokyo assured me this was impossible but the local authorities took a more favorable view & thanks to their good services complete success has crowned our efforts. I have seen, photographed, & collected ample material of every species & variety of conifer known from Formosa. But it must be confessed that some of these Formosan conifers have exacted severe toll in time, money & energy & at the moment of writing I am leg weary & tired.... When last I wrote I mentioned that my next trip had for its principal object Cunninghamia konishii [another member of the cypress family]. Bad weather hampered things but I got him & photographs also. I then switched off to another district & got the Libocedrus [an endemic incense cedar, now recognized as Calocedrus formosana] which now is found only on steep ridges & cliffs almost inaccessible. So difficult is the country that it was not possible to obtain photographs of the whole tree but only sections.... I was back here on April 4th & left the next morning to collect the last remaining species--the For mosan Pseudotsuga [an endemic Douglas fir, now recognized as P. sinensis var. wilsoniana].... On the morning of the fourth day we found our tree but all our efforts to find more failed. Dense fog came on & photography was out of question. The tree was a large one, fully 90 ft. tall & 12 ft. in girth of trunk which divided into three stems.... We then descended some five miles to our lodgings--a police hut--hoping that the next morning would be clear so that we might return & photograph the tree. It rained during the night but morning broke gloriously fine & we got back to the tree by 9 a.m. It stood badly for photography & we were nearly three hours cutting (or rather hacking for our tools were poor) away surrounding trees before a satisfactory picture could be taken. However, fortune favored us but scarcely had we finished when down came the mists blotting out everything. The task accomplished we packed up & returned by the way we came. APRIL 14, 1918 | TAIPEH, FORMOSA Formosa is a land wherein it is quite impossible to travel off the beaten track without official sanction & assistance. To us everything has been open & every wish, expressed or implied, viewed favorably. The director of the forestry experimental station, Mr. R. Kanehira, is a very exceptional man full of energy, enthusiasm & good will, & associated with him are at least two very competent collectors. Kanehira arranged all our trips & accompanied us on most of them. We got to know (he speaks English perfectly) one another pretty well & I hope to our mutual advantage. In fact, whilst the tangible results of our trip are considerable no less important in my opinion is the relationship I have established between the Arnold Arboretum & Kanehira & his associates.... Formosa is certainly a rich & beautiful island & its forest wealth is very great. To have visited the island is a privilege I greatly appreciate & I shall carry away with me none but the pleasantest of recollections. With cordial regards & best wishes, I am, dear Professor Sargent, Faithfully & sincerely yours, E.H. Wilson "},{"has_event_date":0,"type":"arnoldia","title":"Hurried Journey: Botany by Rail","article_sequence":4,"start_page":34,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25655","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060b76d.jpg","volume":76,"issue_number":3,"year":2019,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"DAMERY, J. 2019. HURRIED JOURNEY: BOTANY BY RAIL. ARNOLDIA, 76(3): 34?39 Hurried Journey: Botany by Rail Jonathan Damery I f a pin were dropped in the center of a topographic map of Nevada, it would land amidst a series of low mountain ranges, running roughly north and south. The ranges ripple towards the eastern border of the state, forming an arrangement that looks like a furrowed brow. In 1878, Charles Sprague Sargent, the first director of the Arnold Arboretum, embarked for these arid mountains in what would be the first non-local plant collecting expedition by an Arboretum staff member. Sargent found forests within this area that appeared \"scanty and stunted.\" He counted only seven tree species, of which the single-leaf pinyon (Pinus monophyla) and Utah juniper (Juniperus osteosperma, then considered J. californica var. utahensis) were the most abundant. Despite the limited diversity, Sargent was impressed with the trees for their resilience and age. Some, he estimated, were eight hundred years old, if not older. Sargent began this botanical reconnaissance near the town of Eureka, a silver-mining community located roughly in the center of the state. In 1869, the town consisted of one or two cabins, but by the time Sargent arrived, nine years later, it had grown into the second largest town in the state, with a population, according to boosters, that neared seven thousand. The town boasted a new brick hotel, an opera house that could seat five hundred, two banks, four churches, three newspapers, and, most importantly, sixteen furnaces for smelting silver ore. All of this--along with Sargent's arrival--was facilitated with a narrow-gauge railroad, completed in 1875, which connected Eureka with the town of Palisade, about eighty-five miles to the north. Those tracks, in turn, were made practical by the Pacific Railroad, completed in 1869, which carved its way through Palisade. The Pacific Railroad--composed of the Central Pacific to the west and the Union Pacific to the east--was the first railroad to span the Rocky Mountains and the Great Plains, connecting San Francisco with Omaha and cities beyond. To Sargent, railroad transportation would have seemed ordinary. After all, his father, a banking president named Ignatius Sargent, had been on the board of directors for several New England railroad companies since 1849, and in 1880, Charles would assume his father's membership on one of these boards--the Boston and Albany Railroad--and would continue in that capacity through 1900. Railroads were in the family. Yet when Sargent headed for Nevada, the Pacific Railroad was less than a decade old, and the railroad was just beginning to redefine botanical possibilities in the western United States. Sargent, himself, described his expedition as a \"hurried journey,\" suggesting how remote landscapes had been rendered newly accessible. Unlike botanical explorations that occurred in Nevada before the ceremonial golden spike was driven on May 10, 1869--the date when transcontinental rail passage was inaugurated--Sargent's field research could be conducted in the matter of two weeks (rather than months or years), with the subsequent research publication written in the comfort of Boston and Brookline. By Horse and by Foot In 1955, Susan Delano McKelvey, an Arboretum botanist, published an eleven-hundredpage tome on early botany of the western United States, titled Botanical Exploration of the Trans-Mississippi West: 1790?1850. According to McKelvey, an Englishman named Joseph Burke was one of the first scientifically trained botanist to make observations in central Nevada. Burke spent thirty-eight months in the western United States, beginning in the spring of 1844, and he crossed Nevada in the summer of 1846. His account of the Nevada landscape provides scant details, however, because when that portion of the expedition ended at Fort Walla Walla, in southern Washington, he received two overdue letters from William Jackson Hooker, the director of the Royal ARNOLD ARBORETUM ARCHIVES Botany by Rail 35 A map from Susan Delano McKelvey's Botanical Exploration of the Trans-Mississippi West: 1790?1850 shows the basin-and-range topography of central Nevada. The botanist Joseph Burke (Bu) passed through Nevada in 1846, partially sharing a route used by John Charles Fr?mont (F3) in 1845. Botanic Gardens, Kew, who was the primary sponsor for the trip. The second letter informed Burke that his funding had been halted due to Hooker's dissatisfaction with the amount of collections Burke had provided. Burke defended his record in a long response letter, noting several shipments of seed--the most recent of which had been sent \"across the mountains by the express\" and herbarium specimens that had been sent for a ship in Vancouver. Burke then resigned from the expedition. \"I think, Sir William, it is a very hard case if a collector is sent from the Royal Botanic Gardens to a country where he cannot send his collections by any means by the time mentioned in your letters,\" he wrote. \"I trust, Sir William, you will forgive my retiring from the service without waiting an answer, as it would be two years or upwards before I could receive one.\" It would, in fact, take fourteen months for his letter to arrive on Hooker's desk. So, Burke's estimate was realistic, and without the guarantee of money and supplies in the meantime, his explorations could not continue. He returned home. McKelvey, for her part, felt that Hooker was unfair to Burke, noting the physical rigor associated with backcountry botanical expeditions, where botanists were responsible for travelling with packages of seeds and herbarium specimens--not to mention food and supplies--for weeks if not months on end. \"To work one's way thus encumbered through a pathless wilderness of swamps, undergrowth or fallen timber, up and down ravines, across creeks and rivers, in fair weather or in, veritably, foul or to traverse for days on end waterless deserts in horrible heat and permeating dust, was exhausting work, and the collector was not chosen because he was qualified as a Paul Bunyan,\" McKelvey writes. She goes on to narrate the evening routine botanists were generally obligated to undertake: stopping for camp, building a fire to 36 Arnoldia 76\/3 ? February 2019 prepare food and stay warm (even in the desert), and then arranging the daily collections of plant clippings between layers of paper and pressing them tight. Often, too, given that the papers used for herbarium specimens were prone to become damp or wet throughout the course of a trip, the botanist would need to regularly redo older specimens, transferring them to drier papers, in order to prevent mildew. Plant collecting was (and still is) physically demanding. These routines would have certainly applied to Burke, although it is unclear how many botanical collections Burke made in Nevada. He travelled across the state with a group of settlers that were following a newly blazed trail for Oregon's Willamette Valley. The team consisted of twenty-four individuals and several wagons, and it took nearly seven weeks for them to pass between Fort Hall, on the Oregon Trail, and the Willamette. Burke wrote little about Nevada, but he noted that when the team passed through the northwestern corner of the state, a landscape now known as the Black Rock Desert, it was the \"most miserable volcanic region, with many boiling springs.\" He recorded nothing of botanical interest until spotting an expanse of California poppy (Eschscholzia californica), which decorated a recently burned river valley with papery orange flowers, in southwestern Oregon. The poppy was \"a very shy fruiter,\" he wrote, as was the golden chinquapin (Castanopsis chrysophylla) that he encountered several days later. When they arrived at the Willamette farmstead where the leader of the wagon train lived, the whole team heaved with exhaustion, horses and humans alike. Burke rested three days and then continued to Oregon City--south of Portland. His horses \"nearly drowned\" while swimming a creek on the way (presumably soaking any herbarium specimens that he had collected), and it would take him another two weeks to reach Fort Walla Walla, where his resignation letter was ultimately penned. Over the three decades that separated Burke from Sargent, other botanists passed through northern and central Nevada, and the most detailed observations were rendered by Sereno Watson, who would later become the curator of the Gray Herbarium and Library at Harvard. Watson embarked, in 1868 and 1869, as the lead botanist on two of six field seasons by a geological team surveying the fortieth parallel between California and the Great Plains. Watson's first season focused primarily on central Nevada, the second on Utah--almost entirely within the self-contained watershed of the Great Basin. The region was of interest for the survey (which had begun in 1867) because no accurate maps existed and because the federal government was intent on cataloguing the natural resources along the projected path of the Pacific Railroad. Watson began at Carson City, Nevada, in April 1868, moving east on an indirect path. The purpose of the survey was thoroughness rather than speed, and the team spent a full six weeks working from a basecamp at Fort Ruby, about seventy miles northeast of the prospecting encampment at Eureka (of which Watson makes no mention). From Fort Ruby, explorations were made in the surrounding mountain ranges and valleys. Watson observed several locations where relatively sizeable conifers could be found, including limber pine (Pinus flexilis), growing in the East Humboldt Mountains, with individuals sometimes (though rarely) reaching fifty feet high. Although Watson documented his findings in incredible detail, he wrote little about the comforts or difficulties of travelling with the sur vey team, and he said nothing about the logistics of offloading herbarium specimens for shipment. Nevertheless, had his months in Nevada occurred even one year later, the realities of the railroad would have begun to reshape these considerations. By 1868, railroad workers had already begun to lay tracks across Nevada, and in 1869, these tracks were operational. Therefore, Watson's study marked an important moment: not only had it resulted in the most detailed account of the flora of central Nevada published to date but it also represented the final botanical study in the region before the landscape was bound into the national infrastructure of steel tracks and steam locomotion. Geologists on the survey would subsequently comment about strata and fossils observed at railroad cuts, indicating how the presence of the railroad became ingrained in the researchers' world. Botany by Rail 37 Botanical Space Given that Watson and Sargent would become Harvard colleagues, the men must have conferred about the flora and landscape of central Nevada while Sargent was making travel preparation in 1878. Yet Sargent also saw his trip as a follow-up to an expedition the previous summer by Asa Gray--the preeminent Harvard botanist--and Joseph Dalton Hooker, the English botanist who had assumed his father's role as director of the Royal Botanic Gardens, Kew. When Sargent returned from his trip, he sent Hooker a letter recounting his findings in detail, noting that he expected Hooker would remember the Palisade station on the railroad. Sargent continued south to Eureka, whereas Hooker and Gray continued riding the Pacific Railroad to Carson City. Yet the implications of Hooker's presence in this region is significant, given that thirty-one years before, Joseph Burke was passing through this exact same stretch-- then remote and without a defined wagon route--under the direction of Hooker's father. The son, acting in the same official capacity as director of Kew, was making a passage that his father had commissioned another to make. McKelvey stresses the power dynamics that were often at play between collectors and the individuals who sponsored their trips. She notes that few of the botanists considered in her book--individuals working in the western United States before 1850--were engaging in their own independent research. \"By far the greater number went at the behest of professional botanists living in proximity to the essentials of herbaria and libraries, and in distinction to their emissaries, amid safe and comfortable surroundings,\" McKelvey writes. \"The backers of the scheme--often called `closet botanists' for the reason that, working in offices, they may never have seen the living plants which they described--were engaged for the most part in descriptive botany, writing botanical papers or compiling floras of small or large scope.\" While Joseph Hooker began his early career with an expedition to Antarctica (and the surrounding islands) and then another to India, those two expeditions collectively required more than seven years abroad. The fact, therefore, that Hooker could now spend scarcely three months travelling from the Atlantic Coast to the Pacific Coast of the United States and back was a radical convenience. Instead of sending an explorer with youthful enthusiasm and resilience-- someone like Burke--Hooker himself could go, even as a sixty-year-old and even as the director of a major botanical institution. For Gray, this was a second trip on the Pacific Railroad; the first was in 1872. Gray's wife, Jane Loring Gray, accompanied him on both trips, and she would later recall him racing into the landscape at short station stops, collecting whatever he could find. This caused considerable intrigue for fellow passengers, who then gathered around to watch Gray prepare his herbarium specimens. Eventually others began to collect plants as well, bringing them to Gray for identification and causing exasperation for the conductor. It took them a day to cross Nevada, where Gray noted the snaking green vegetation along the Humboldt River. In a letter to his friend Richard William Church, Gray described the whole experience with exceptional enthusiasm. \"There were fatigues and small discomforts, of course, but these are all forgotten long ago, and the whole transit dwells in memory as one continual and delightful piece of pleasant, novel, ever-varied, and instructive sightseeing,\" he wrote. \"Of course, the identifying at sight, as we flew by, of flowers new to me in the living state, and the snatching at halts, and the physical features of districts which I had always been interested in, and knew much about but had never seen, all gave me occupation and continual pleasure.\" In this way, the Pacific Railroad was beginning to reshape botanical space in the western United States. By the time the Grays made their rail passage in 1872, five hundred miles through the Great Basin no longer meant the same thing that it had with Watson's expedition a mere four years before, let alone more than two decades before with Burke. While botanical explorations in the region could still be physical and immersive, the work was conducted with two steel lifelines to urban centers. Herbarium specimens no longer needed to be transported for weeks or months before reaching a shipping location. While Sargent and other leading botanists would continue to enlist field 38 Arnoldia 76\/3 ? February 2019 collectors to work in the western United States, the power and money associated with collecting along these railroad axes had been forever transformed. Botanical Limits COURTESY OF THE ARCHIVES OF THE GRAY HERBARIUM Certainly, the Pacific Railroad did not uniformly influence botanical space in the western United States, and in summer of 1883, Sargent would participate in a geological survey associated with the installation of the Northern Pacific Railroad that connected Tacoma, Washington, with St. Paul, Minnesota. Although he was gone less than two months, that expedition was rife with peril, including two instances where pack animals slipped and fell precipitously. (In the second case, the horse fell fifteen hundred feet, carrying Sargent's plant collections and the team's guns.) Yet Sargent, like others, quickly understood that it wasn't just botanists that would be benefit from this reconfiguration of space along the railroads. The power to study these landscapes came with the simultaneous power to exploit the resources found therein. Both processes could be conducted at an unprecedented rate. In 1878, after Sargent arrived in Eureka, he obtained a wagon and continued southwest for about seventy-five miles, exploring the Monitor Range, which reaches points well over ten thousand feet above sea level. He then continued to Carson City, from which he proceeded into California. During his two weeks in Nevada, he collected a considerable amount of seed, which he planned to introduce into garden cultivation. Meanwhile, he became increasingly attuned to the risks facing these unassuming and hardscrabble forests. Wood of the Utah juniper (Juniperus osteosperma) was widely harvested for cheap fuel, given that it was the only tree Harvard botanist Asa Gray rests at La Veta Pass, Colorado, holding an herbarium press (foreground), with Kew's Joseph Hooker at his side (seated at left). Jane Loring Gray wears a light-colored coat at the table behind. This 1877 expedition continued to California using the Pacific Railroad. Botany by Rail 39 found abundantly at lower elevations. (Its wood even powered the steam locomotive on the Eureka and Palisade Railroad.) Other tree species were harvested for lumber, charcoal, and even bearings for machinery. Most striking, however, were his observations of the Great Basin bristlecone pine (Pinus longaeva, then considered P. balfouriana). He found several specimens, growing between fifteen and thirty feet tall, on a mountain near Eureka. \"Formerly the whole summit of this mountain was very generally covered with this species,\" he wrote, \"but with few exceptions the trees have all been cut to supply the mines with timbering, for which purpose the strong and very close-grained, tough wood of this species is preferred to that of any other Nevada tree.\" Sargent didn't estimate the age of these trees or count the tightly packed growth rings, but in California, this species is now known to reach more than five thousand years old. On the same mountain, Sargent observed a curl-leaf mountain mahogany (Cercocarpus ledifolius)-- a small tree in the rose family (Rosaceae)--and he suggested that plant was least 890 years old, if not much older. \"It is perhaps permissible to suppose that the seed which produced this little tree had already germinated when the oldest living Sequoia on the continent was still a vigorous sapling with its bi-centennial anniversary still before it,\" Sargent wrote. Sargent suspected that someone travelling across the Great Basin on the Pacific Railroad would perceive a landscape that was essentially \"destitute of trees,\" much like the prairies to the east. Yet he came to recognize the \"immense value\" of the forests, no matter how diminutive. \"It will have been seen that the forests of Nevada, consisting of a few species adapted to struggle with adverse conditions of soil and climate, are of immense age, and that the dwarfed and scattered individuals which compose them reach maturity only after centuries of exceedingly slow growth,\" he wrote. \"On this account, and because, if once destroyed, the want of moisture will forever prevent their restoration, either naturally or by the hand of man, public attention should be turned to the importance of preserving, before it is too late, some portions of these forests.\" He proposed that the federal government should step in to preserve the remaining woodlands, warning that \"terrible destruction\" would occur otherwise. (About three decades later, the Humbolt-Toiyabe National Forest was established, protecting vast swaths of these noncontiguous mountain forests.) In this sense, Sargent's railroad-powered expedition allowed him to articulate the finite limits of botanical space. Forests that were once remote and practically inaccessible for a Bostonian like Sargent were now mere days away, and their future, as a result, seemed ever more precarious. References Ambrose, S.E. 2001. Nothing like it in the world: The men who built the transcontinental railroad, 1863?1869. New York: Touchstone. Burke, J. 1846. Burke to W.J. Hooker, 17 October 1878. Director's correspondence, Library and Archives at Royal Botanic Garden, Kew. Gray, J.L. (Ed.). 1894. Letters of Asa Gray (Vol 2.). Boston and New York: Houghton, Mifflin, and Co. Lambert Molinelli and Co. 1879. Eureka and its resources: A complete history of Eureka County, Nevada. San Francisco: H. Keller and Co. McKelvey, S.D. 1955. Botanical Exploration of the TransMississippi West: 1790?1850. Jamaica Plain, MA: Arnold Arboretum of Harvard University. Sargent, C.S. 1877. Sargent to J. Hooker, 21 October 1878. Director's correspondence, Library and Archives at Royal Botanic Garden, Kew. Sargent, C.S. 1879. The forests of central Nevada, with some remarks on those of the adjacent regions. The American Journal of Science and Arts, 3: 417?426. Sutton, S.B. 1870. Charles Sprague Sargent and the Arnold Arboretum. Cambridge, MA: Harvard University Press. Watson, S. 1871. Botany. In C. King (Ed.) Report of the geological exploration of the fortieth parallel (Vol. 5). Washington, DC: Government Printing Office. White, R. 2011. Railroaded: The transcontinentals and the making of modern America. New York: W. W. Norton and Co. Williams, R.L. 2003. A region of astonishing beauty: The botanical exploration of the Rocky Mountains. Lanham, MD: Robert Rinehart Publishers. Jonathan Damery is the associate editor of Arnoldia. "},{"has_event_date":0,"type":"arnoldia","title":"Sax Pine: A Hybrid Left Behind","article_sequence":5,"start_page":40,"end_page":41,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25657","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060bb6b.jpg","volume":76,"issue_number":3,"year":2019,"series":null,"season":null,"authors":"Rubinstein, Jared","article_content":"RUBINSTEIN, J. 2019. SAX PINE: A HYBRID LEFT BEHIND. ARNOLDIA, 76(3): 40 Sax Pine: A Hybrid Left Behind Jared Rubinstein L ast fall, gray fog streamed down the hillside by the Hunnewell Building, enveloping the magnolias and eastern white pines (Pinus strobus) that dominate the area. The display emanated from an art installation, Fog x FLO, by Fujiko Nakaya, and when viewing the fog from atop the hill, as signs encouraged visitors to do, a few pines stuck out from the others. With bluish needles and scaly bark, these trees didn't look quite like their neighbors, nor did they look quite like any other species of pine. That's because these trees (accession 266-46) are hybrids. Although the Arnold Arboretum is best known for its wild-collected plants, most identified to a single species, we also have a significant collection of hybrid plants, including many that were bred and developed here. Karl Sax, a professor of botany at the Bussey Institute and later director of the Arboretum, created some of the Arboretum's best-known hybrids, including Forsythia `Meadowlark' and Magnolia ? loebneri `Merrill', which both can be found growing in the Arboretum and around the world. But Sax didn't only work with flowering trees or shrubs--he also dabbled with conifers. In the early 1940s, Karl Sax went on a bit of a pine hybridization kick. Crossing different plant species can be tedious: Pines are wind pollinated, so Sax covered the female cones of one pine species with a bag to prevent natural pollination from pollen blowing around in the wind. When the time was right, he removed the bags and introduced pollen collected from male cones of a different pine species to the female cones. Once the hybrid seeds had developed within the cone, Sax removed and planted the seeds in the nursery at the Bussey Institute. Sax mixed and matched pines from all over the world--New England pines with Himalayan pines, European pines with Japanese pines, West Coast pines with East Coast pines--all with an eye towards producing something new with a high economic or ornamental value. The hybrid pines behind the Hunnewell Building are crosses between Pinus monticola, the western white pine, and P. parviflora var. himekomatsu, the southern variety of the Japanese white pine. The combination shows just how well hybridization can capture traits from each parent. The needles, in fascicles of five, maintain the long, soft appearance of P. monticola but gain a glaucous, blueish-gray color from P. parviflora. The hybrids seem to get their height from P. monticola, especially accession 266-46*B, which soars to almost 75 feet (23 meters), already much higher than even the oldest Japanese white pines at the Arboretum. And the bark, normally thin and smooth on P. parviflora and rough and flaky on P. monticola, forms elegant plates that are divided into scales--a sort of middle ground between the two parents. When evaluating hybrids, one usually looks for hybrid vigor, or traits that give a hybrid an advantage over its parents, like a better form or a higher tolerance to adverse environmental conditions. While these particular hybrids do appear to be vigorous growers and have an unusual mix of features, they never managed to achieve the fame found by some of Sax's other hybrids, like Prunus `Hally Jolivette' or Malus `Mary Potter'. The beauty of these hybrid pines is perhaps a more subtle one, and they just weren't flashy enough to make it big in the horticultural industry of the 1940s. Unlike other pine hybrids Sax tried out, such as Pinus ? hunnewellii or Pinus ? schwerinii, these hybrids were never given a nothospecies designation--that is, a Latin name specific to that hybrid. What's more, these hybrids do not appear in horticultural catalogs or seem common in other arboreta. Far from diminishing their value, however, this lack of fame makes these hybrids all the more special to the Arboretum. It's possible that the five plants growing here are the sole representatives of this hybrid in cultivation. More than anything, these hybrids highlight the importance of experimentation and of following curiosity to wherever it may lead. Their longevity and beauty remind us that even hybrids that don't \"make it\" deserve another look. Jared Rubinstein is the living collections fellow at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25746","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d160bb6f.jpg","title":"2019-76-3","volume":76,"issue_number":3,"year":2019,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Behind the Scenes on Naming a New Hemlock Species","article_sequence":1,"start_page":2,"end_page":9,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25649","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060ab28.jpg","volume":76,"issue_number":2,"year":2018,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Behind the Scenes on Naming a New Hemlock Species Peter Del Tredici I n my thirty-five years working at the Arnold Arboretum, I have only once been involved in naming a new species--a hemlock tree (Tsuga ulleungensis) native to the volcanic island of Ulleungdo in the Sea of Japan. This South Korean island lies 80 miles east of the mainland and 176 miles from the west coast of Japan. It's a small island--twenty-eight square miles--and rises to an elevation of thirty-two hundred feet. Geological records indicate that Ulleungdo emerged from the ocean floor some 2.7 million years ago and that it has been vegetated for the past 1.7 million years. The story of how I got involved with naming this new hemlock tree is complicated, but it bears telling for what it reveals about the process of doing basic botanical research and about how welldocumented collections of plants--leavened with a bit of serendipity--can contribute to research on evolution and biogeography. Although I was not aware of it at the time, my involvement with the plant actually began with the arrival of hemlock woolly adelgid (Adelges tsugae) at the Arboretum in 1997. This invasive sucking insect is native to Japan and was discovered in the eastern United States in the 1950s. It took about forty years for the bug to spread from the site of its original introduction in Richmond, Virginia, to the Arboretum--its relentless northward spread driven by warmer winters induced by climate change. When it finally arrived in 1997, it struck with a vengeance, attacking over a thousand trees located on the twenty-two acres of Hemlock Hill and scattered throughout the rest of the grounds. While the insect certainly presented serious problems for the Arboretum, it posed a far bigger threat to the wild-growing hemlocks of southern New England. In response to the arrival of hemlock woolly adelgid in the Northeast, biologists adopted a multipronged approach to try controlling the pest: Plant physiologists evaluated various chemical controls. Entomologists began searching for Asian insects that might be used as potential biocontrol. And horticulturists began evaluating other hemlock species for resistance to the bug. Given the Arboretum's extensive collection of hemlock species, which is nationally accredited through the American Public Gardens Association's Plant Collections Network, a number of researchers began coming here in hopes of finding something--anything-- that might shed light on the problem and contribute to a solution. Among these researchers was a young graduate student in entomology at Yale University, Nathan Havill, who was also working at the United States Forest Service's Northern Research Station in Hamden, Connecticut, trying to unravel the insect's complex life cycle and assess its genetic diversity across the region. As luck would have it, Michael Donoghue, a graduate and former professor at Harvard, was on Nathan's dissertation committee at Yale and suggested that just studying the insect was not enough and that Nathan should also to do genetic work on hemlock trees. Now anyone who has been a graduate student knows how annoying it can be when a committee member adds more work to a thesis project, but Nathan recognized a good idea when he saw one and agreed to investigate the genetics of the Arboretum's hemlocks in order to create a phylogenetic tree describing their relatedness. For help, he recruited Chris Campbell, a professor at the University of Maine and another Harvard alum, to help with the project because one of Chris's former students, Tom Vining, had produced DNA sequence data on Tsuga and developed a rough phylogeny of the genus in 1999. I first met Nathan in the spring of 2003, when he came to collect hemlock samples at the Arboretum. He was also looking for adelgid galls on the bug's alternate host, the Japanese tigertail spruce (Picea torano). It wasn't until the following year, having completed his genetic analysis, that Nathan shared the preliminary Tsuga ulleungensis 3 Tsuga sieboldii. This was a logical decision on his part given close morphological similarities with the southern Japanese hemlock and the fact that there were no hemlocks on the Korean peninsula or adjacent parts of China. Nakai's classification held up until 2008 when Nathan published his finding that the Ulleungdo hemlock was genetically distinct from the southern Japanese hemlock. Although Nathan indicated that more detailed analysis was required, his results suggested the taxon was more closely allied with the northern Japanese hemlock (T. diversifolia). A few months before this paper was published, I met up with Nathan at the Fourth Symposium on the Hemlock Woolly Adelgid, held in Hartford, Connecticut, in February 2008. I asked him about the Ulleungdo hemlock, and he told me that his research was inconclusive as to whether or not it was a new species. I was surprised to hear this and immediately asked him if he would have a problem JONATHAN DAMERY results of his research with me and casually mentioned that one of our accessions--labelled as southern Japanese hemlock (Tsuga sieboldii), from Ulleungdo, South Korea--did not line up with the Arboretum's other specimens under the same name, which was weird. Our two plants from Ulleungdo (accession 1251-83*A and B) had arrived in 1983 as wild-collected seed obtained by the Chollipo Arboretum, an institution located outside of Seoul, and no one prior to Nathan had questioned their identity. Despite its small size, Ulleungdo is famous in botanical circles for harboring over thirty species found nowhere else in the world. Ernest Henry Wilson visited the island in the late spring of 1917, guided by the botanist Takenoshin Nakai who was describing the island's plants on behalf of the Japanese government, which had taken over the kingdom of Korea in 1910. Wilson collected herbarium specimens of the hemlocks that he found there and, following Nakai's lead, referred to them as Two anomalous hemlocks on Bussey Hill became the catalyst for naming a new hemlock species (Tsuga ulleungensis), which is endemic to Ulleungdo, South Korea. Accession 1251-83*B is shown here, overlooking the oak collection. ARNOLD ARBORETUM AND GIS COMMUNITY 4 Arnoldia 76\/2 ? November 2018 Ulleungdo is a small volcanic island, ecologically isolated within the Sea of Japan. Botanists have discovered more than thirty endemic plant species or subspecies on the island. Collecting locations by Peter Del Tredici and his colleagues are marked. with me continuing his work on the taxonomic status of Ulleungdo hemlock. That would be fine, he said, because he was now working full time for the Forest Service on the genetics of the hemlock woolly adelgid and had no time for continued research on the trees. He also suggested that I collaborate with Ashley Lamb, an entomologist at Virginia Tech, who was going to southern Japan in the spring to study a potential hemlock woolly adelgid biocontrol insect, a small beetle known as Laracobius osakensis. I located Ashley at the conference later that day, and in no time flat, we agreed to collaborate on an Arboretum-funded trip to Ulleungdo. We would collect hemlock leaf samples and look for adelgids that Nathan could sequence for his research. Not even three months later, on May 9, Ashley and I met in Osaka, Japan, and immediately caught a flight to Seoul. There, we made contact with Nam Sook Lee, a professor at Ehwa Womans University, and her colleague Sung Hee Yeau. I had met Nam Sook several years earlier when she was visiting herbaria in the eastern United States--including Harvard--and we had kept in touch. When I began developing my plans to go to Ulleungdo, Nam Sook generously agreed to host the expedition, which meant making all of the travel arrangements and procuring the permits needed to collect research samples on the island. The four of us left early the next morning on the three-hour drive from Seoul to Donghae, the point of departure for the RAKI MAN Tsuga ulleungensis 5 ferry to Ulleungdo. It was a clear day, but the seas were rough. Despite my scopolamine patch, I suffered mightily from sea sickness during the seemingly endless journey, as did most of the other passengers on the ship. As far as I could tell, only Ashley, sleeping with her head down on a table the whole time, emerged unscathed. Needless to say, I was relieved when our ship finally landed at Jeodong, one of the island's two main cities, where we were met by Suk Su Lee, Dodong, the primary port city for Ulleungdo, is located south of Jeodong. In the forewho worked for the local ground, Sorbus ulleungensis, an endemic mountain ash, can be seen flowering. forestry department. lock trees that he had collected from the forest (Later, from 2011 to 2015, he served as chief and planted at the entrance. They were growof forestry for Ulleung County, which also ing extremely well--about twenty-five feet tall includes forty-three smaller islands). Suk Su and very full--and all three were loaded with quickly found a taxi and took us to our hotel, developing cones. where I promptly passed out for two hours. He We were up early the next day and took a taxi met us later for lunch (which I skipped, still across the island to Taeha Ryeong, a special feeling queasy), and then he drove us up into reserve established in 1962 to protect some of the hills north of Jeodong, nearly one thousand the largest hemlocks, beeches, and Japanese feet in elevation, where I got my first look at the native forest with Camellia japonica white pines on the island. The slopes were in bloom in the understory, along with the steep, and the hemlocks were especially large Ulleungdo beech (Fagus crenata var. multinerin trunk diameter--up to 25.5 inches (65 cenvis), which many taxonomists treat as a sepatimeters). I managed to collect leaf samples rate species (F. multinervis). We also observed from about twenty trees that had branches low two varieties of maple endemic to Ulleungdo enough to reach; all were growing around fif(Acer mono ssp. okamotoanum and A. pseuteen hundred feet in elevation. From there, we dosieboldianum ssp. takesimense), the Japahiked down the mountain through a patch of nese white pine (Pinus parviflora), and most woods that harbored an unbelievably rich herimportantly, the Ulleungdo hemlock, the target baceous understory. We ended up at a clearing of our expedition. where our taxi driver was supposed to meet us, On the second hemlock tree we examined, but since he wasn't there, we took the opporAshley found evidence of the adelgid and immetunity to do more collecting. Suk Su, at some diately collected samples for Nathan. We stayed risk to life and limb, climbed a tall hemlock in the area for a couple of hours, collecting more and managed to collect a branch with both male samples of the adelgid as well as herbarium and female cones on it, which we had been specimens and leaf samples of the hemlocks. unable to obtain at our earlier location because On the way down, the sun was setting, and we the trees were too tall. He climbed down from the tree with the precious specimen just as our stopped at the elementary school in Jeodong, where Suk Su proudly showed us three hemcab showed up. PETER DEL TREDICI 6 Arnoldia 76\/2 ? November 2018 Collaborator Suk Su Lee collected Tsuga ulleungensis from the wild and planted three specimens near the elementary school in Jeodong. After Peter Del Tredici visited in the spring of 2008, Suk Su sent seed from these plants to the Arboretum, where they were propagated and planted in the landscape. From Taeha Ryeong, we drove halfway around the island to the Nari Basin, a volcanic caldera left after an eruption that occurred about ten thousand years ago. This is now the only place on the island with relatively flat ground, so the locals have taken advantage of this fact by establishing agricultural fields devoted to the cultivation of local medicinal plants, such as Codonopsis lanceolata, a vining member of the bellflower family (Campanulaceae). The landscape was beautiful, ringed by mountains, with specimens of the Ulleungdo hemlock in the surrounding forest and Sorbus ulleungensis, an endemic mountain ash, growing at the edges of the fields. (Incidentally, the mountain ash was then considered to be S. commixta-- a species that ranges through northern Japan and the islands of eastern Russia--illustrating another case where speciation on Ulleungdo was long unrecognized.) We had lunch at a vegetarian restaurant that served the traditional bibimbap dish filled with medicinal plants cultivated in the caldera and wild plants from the surrounding mountains--super healthy we were told--which was unlike anything I've tasted before or since. The caldera was a jumping-off point for hiking to the highest peak on the island, but it was too late in the day for us to make the trek, so we headed back down the mountain to our hotel. We were greeted with bad news that a storm was moving in and that we would have to leave the island a day earlier than planned or risk getting stuck and missing two lectures I had scheduled in Seoul. The coup de grace was that there were no seats left on the boat leaving at five o'clock the following afternoon, so our only choice was to take a ferry at five the next morning. In the absence of any alternative, I packed up the specimens and set my alarm clock. It was frustrating to have come this far only to have the trip cut short before I could finish exploring the island. The good news was that I had collected leaf samples from PETER DEL TREDICI ARNOLD ARBORETUM ARCHIVES Tsuga ulleungensis 7 Ernest Henry Wilson photographed a stand of hemlocks--now Tsuga ulleungensis--on Ulleungdo in 1917, estimating heights ranging to 75 feet (23 meters). Peter Del Tredici observed similarly impressive specimens at the Taeha Ryeong Reserve in 2008, including this tree, which measured 98 feet (30 meters) tall. thirty-one plants in five different locations, which was enough for a thorough genetic analysis of the Ulleungdo hemlock population. After returning to the Arboretum from Korea, I continued making observations on the leaf and cone morphology of our two specimens of the Ulleungdo hemlock, but I failed to make progress on the genetic analysis of the leaf samples. The breakthrough finally came two years later, in August 2010, at the Botanical Society of America meetings in Providence, Rhode Island. I met with Chris Campbell, the professor from the University of Maine who had collaborated with Nathan on hemlock phylogeny, and his graduate student Garth Holman. After a brief discussion, we all agreed that Garth should include a chapter on the Ulleungdo hemlock in his dissertation on conifer genetics. His research would be based on the Arboretum's trees and on the DNA samples I had collected from Ulleungdo. It was with great joy and relief that I turned my samples and herbarium specimens over to Garth later that fall--at last they were going to someone who would put them to good use. For the next four years, Garth labored in the lab on the genetic analysis of various hemlock species, while also assessing their morphological variation. At the same time, I continued with my phenological studies at the Arboretum, which showed that the Ulleungdo hemlock consistently leafed out a few days before Tsuga diversifolia and more than a month before T. sieboldii. With the help of Arboretum staff member Kevin Block, I also determined that the Ulleungdo hemlock was tolerant but not immune to damage by the hemlock woolly adelgid. Garth completed his dissertation in September 2014 with a preliminary description of the Ulleungdo hemlock as a new species: Tsuga ulleungensis. Going from a dissertation to an actual publication in a scientific journal, however, is not always a quick process. In this case, PETER DEL TREDICI 8 Arnoldia 76\/2 ? November 2018 Comparison of four Asian hemlocks growing at the Arnold Arboretum. On the left, the branches show simultaneous phenological conditions on May 23, 2012. The central images show the attachment point for the seed cones, where differences in the shape of cone-scale bracts are most evident--providing a subtle but important diagnostic characteristic for the species. Tsuga ulleungensis 9 it took Garth another two years to expand and improve the research he had done for his degree, before submitting for publication to Systematic Botany in 2017. The final results, which were published in December of that year, painted a complicated picture of the Ulleungdo hemlock's ancestry: its chloroplast DNA indicated a closer relationship to T. diversifolia than to T. sieboldii, while the nuclear DNA was equivocal about the relationships among the three species. The morphological data--its cone and leaf structure--indicated that the Ulleungdo hemlock was more similar to T. sieboldii than T. diversifolia. Taken together, this evidence indicates that T. ulleungensis is distantly related to both Japanese hemlocks but is probably closer to T. diversifolia than to T. sieboldii. The Ulleungdo hemlock is most likely a remnant of a species that was once widespread on the Korean peninsula but disappeared from the mainland as a result of multiple glaciations that took place during the Pleistocene, over the past million or so years. Because of its mild, oceanic climate, Ulleungdo is the only place in Asia where Tsuga ulleungensis survived. The existence of some thirty-three other endemic species or subspecies of plants on Ulleungdo provides further evidence that this isolated landmass has long served as a glacial refugium. To me, the most interesting thing about the Tsuga ulleungensis story is how it illuminates the role that well-documented living collections can play in supporting basic scientific research and conservation. We start with a graduate student sampling the collections and finding a genetic anomaly. Next, we check the records to see where the plant originated, and then, in collaboration with scientists from the host country, we go back to the original location to recollect the plant for more in-depth analysis. Without the Arboretum's well-managed curatorial system, it would be impossible to make any of these connections. In the case of the Ulleungdo hemlock, a plant growing inconspicuously on the grounds of the Arboretum for twenty years unexpectedly provided a key to understanding the complex evolutionary history of a species restricted to a tiny island off the coast of Korea--half a world away. From a research perspective, the Arboretum's collections are a means to an end rather than an end in themselves: they provide easy access to a significant percentage of the world's temperate trees and shrubs and are the perfect jumping-off point for any number of studies that can open the doors to basic evolutionary questions. References Del Tredici, P., and A. Kitajima. 2004. Introduction and cultivation of Chinese hemlock (Tsuga chinensis) and its resistance to hemlock woolly adelgid (Adelges tsugae). Journal of Arboriculture, 30(5): 282?286. Havill, N., C. S. Campbell, T. Vining, B. LePage, R. Bayer, and M. Donoghue. 2008. Phylogeny and biogeography of Tsuga (Pinaceae) inferred from nuclear ribosomal ITS and chloroplast DNA sequence data. Systematic Botany, 33: 478?489. Havill, N. P., M. E. Montgomery, G. Yu, S. Shiyake, and A. Caccone. 2006. Mitochondrial DNA from hemlock woolly adelgid (Hemiptera: Adelgidae) suggests cryptic speciation and pinpoints the source of the introduction to eastern North America. Annals of the Entomological Society of America, 99(2): 195?203. Havill, N. P., and M. E. Montgomery. 2008. The role of arboreta in studying the evolution of host resistance to the hemlock woolly adelgid. Arnoldia, 65(3): 2?9. Holman, G., P. Del Tredici, N. Havill, N. S. Lee, R. Cronn, K. Cushman, S. Mathews, L. Raubeson, and C. S. Campbell. 2017. A new species and introgression in eastern Asian hemlocks ( P i n a c e a e : Ts u g a ) . S y s t e m a t i c B o t a n y, 42(4):1?15. Wilson, E. 1918. The vegetation of Korea. Transactions of the Korea Branch of the Royal Asiatic Society, 9: 1?17. The map in this article was created using Esri, HERE, Garmin, ? OpenStreetMap contributors, and the GIS user community. Sources: Esri, USGS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, lntermap and the GIS user community. Peter Del Tredici is Senior Research Scientist Emeritus at the Arnold Arboretum and the former Director of Living Collections. He is currently teaching in the Department of Urban Studies and Planning at MIT and finalizing the second edition of his book, Wild Urban Plants of the Northeast: A Field Guide. "},{"has_event_date":0,"type":"arnoldia","title":"Dormant Vines, Future Wines","article_sequence":2,"start_page":10,"end_page":17,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25650","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060ab6d.jpg","volume":76,"issue_number":2,"year":2018,"series":null,"season":null,"authors":"Londo, Jason; Kovaleski, Al","article_content":"Dormant Vines, Future Wines Al Kovaleski and Jason Londo W hen you sip an aromatic Riesling with dinner in September, the day may feel noticeably shorter than it did a month before. While fall officially starts with the autumnal equinox, which occurs between September 21 and 24 from year to year, day length continuously decreases after the summer solstice and will continue through the arrival of winter. From August to November, in fact, the days get shorter by almost three minutes per day in Boston. Plants, including the common grapevine (Vitis vinifera), pay close attention to day length from the moment leaves and flowers unfurl in the spring until fruit ripens and leaves drop. Although this internal clock may seem difficult to conceptualize--more esoteric than flavor and mouthfeel, or even pests and diseases--understanding the process by which plants enter and exit dormancy, and how they survive in between, is critical, especially as winegrowers (and other agricultural producers) adapt their production to withstand a changing climate. As grapevines grow, they form buds in the leaf axils. Within these buds, about ten nodes are pre-formed--leaf primordia (baby leaves) and inflorescence primordia (baby flowers). These buds are formed in preparation for the following growing season. In that way, wines produced in 2018 actually started as microscopic inflorescences in the summer of 2017, which overwintered inside buds, formed flowers that bloomed in the spring of 2018, and became fruit that was harvested in the late summer and fall, effectively spending over a year developing on the plant alone. This process doesn't happen only in Vitis vinifera, the species native to the Mediterranean from which most of our wines are produced. Wild grape species like the riverbank grape (V. riparia), which is native to North America, and gloryvine grape (V. coignetiae), which ranges from eastern Russia to Japan, go through the same process, as do the majority of temperate perennial plants. Unlike migratory birds that avoid winter by flying south, plants are stationary. Therefore, plants must endure low temperatures and other unsuitable growing conditions that come with the winter. In these conditions, they lower their metabolism and enter dormancy, much like hibernation for animals. Plants, however, have evolved to start the preparation for winter much before damaging temperatures arrive, without relying on year-to-year weather patterns. Leaves register the decreasing day length (or technically the increasing nighttime), which provides a signal for buds to enter dormancy. Plants then start changing color to create the beautiful spectacle of fall foliage. In this process, grapes relocate nutrients from the leaves into storage tissue in the woody vines, ready to be recycled the following year. Once plants lose their leaves, they no longer track the day length. Instead, grapevines know when to start growing in the spring by tracking how long they have been cold. Much like a person setting an alarm to have eight hours of sleep, buds count the number of hours spent in what are called chilling temperatures, between 32 and 50?F (0 and 10?C). Just as different people need different amounts of sleep to be productive, each grapevine species, and even different genotypes within a single species, have different chill requirements before they are able to come out of dormancy. Once they have accumulated enough chilling hours, buds are able to better respond to warm temperatures and produce spring growth. The chilling requirement is associated with the region where a species originated: species from lower latitudes are accustomed to low chill accumulation because of short winters, while species from higher latitudes have a high chill requirement because of longer winters. Given that no chill accumulation occurs below 32?F (0?C), however, plants growing in cold continental climates, like Minnesota, or much higher latitudes, like northern Canada, expe- ALL PHOTOS BY AL KOVALESKI UNLESS NOTED Grape Dormancy 11 Because species and cultivars of grape (Vitis) can be grown in a wide range of temperate climates, they prove valuable for studying how hardy plants endure cold weather. At left, October snow clings to Vitis vinifera `Cabernet Sauvignon' in Geneva, New York. Other species, like riverbank grape (V. riparia), shown at right, are even better adapted to cold temperatures. rience low chill accumulation. Species from these places, therefore, also tend to have a low chill requirement. Chill requirements prevent plants from resuming growth during midwinter warm spells, which could cause death of both flowers and leaves upon the return of cold weather. The mechanisms that make buds wait for the appropriate and consistent warm weather in the spring are still largely unknown, but solving this mystery is important. As temperatures continue to increase due to climate change, the amount of chilling that plants experience in different areas is changing: lower latitudes are experiencing fewer hours with chilling temperatures, whereas the opposite is true for higher latitudes, like Boston and upstate New York. This trend in northern areas may seem a little counterintuitive, but time that was previously spent in below-freezing temperatures is now rising into the chilling range, above freezing but below 50?F (10?C), causing chill accumulation to rise. When plants fail to accumulate the necessary chilling requirement, they have erratic, reduced, and delayed budbreak. In vineyards and orchards, this means reduced yields. The same is true with native forests, where flowering and corresponding seed production drops. Moreover, shifting phenology could detrimentally impact overlap between flowering and the activity of pollinators for a given species, or there may be competition due to overlapping flowering for species that were previously staggered. In areas where excessive chill is expected, more responsive plants may break buds during midwinter warm spells, when they previously would have known to wait. In agricultural settings, new cultivars and adaptive management practices can 12 Arnoldia 76\/2 ? November 2018 Grapevines, like those in this commercial vineyard in Geneva, New York, gradually prepare for cold temperatures by tracking the increasing length of uninterrupted nighttime in the fall. help overcome these effects in the short term. For forests, however, climate change is happening faster than floristic composition can change. Researchers at the University of Alberta, in Canada, have estimated that forest species are already 80 miles south of their optimal climate niche, and this figure is expected to increase to 190 miles in the 2020s, beyond recorded rates of natural forest migration. Dormancy Fundamentals Vitis is a compelling genus to study dormancy because of its distribution from tropical to subarctic regions. Even the common grapevine (V. vinifera) alone is capable of growing within a wide band of climates, with the majority of production in the Northern Hemisphere spanning latitudes between San Diego and Vancouver (30 to 50? north). In the Southern Hemisphere, the band is even wider, stretching from northern Argentina to southern New Zealand (20 to 50? south), with some production occurring as close to the equator as northeastern Brazil (9? south). Moreover, many of the most popular cultivars like V. vinifera `Pinot Noir' and `Chardonnay' are present in almost all of these areas, which demonstrates the remarkable plasticity of the species. Also, many varieties have been cultivated for a very long time. The first records for `Riesling' date from the fifteenth century-- before Columbus arrived in the Americas--and impressively, seeds of `Rkatsiteli' were found in clay vessels dating to 3,000 BC. `Rkatsiteli' is still grown in Georgia, the eastern European country where the archaeological discoveries occurred, and limited production can be found in the Finger Lakes wine region of New York. This historical continuity provides us with a wealth of records from different regions, providing both temporal and geographic context for understanding the basic requirements for dormancy of grapevines. Measuring the chilling requirement of different grapevine varieties can be very simple, and in fact, similar techniques can be used to study dormancy in most deciduous perennial plants. Grape Dormancy 13 Once leaves have dropped, buds in the vineyard will wait to unfurl until a specified amount of chilling temperatures has been experienced. Cuttings with dormant buds that have experienced different amounts of chill are placed in a warm environment (72?F or 22?C) with sixteen hours of daylight. The chilled buds can either be collected from the field in regular intervals throughout the winter, or collected in late fall or early winter and placed in a cold chamber where they'll be removed after certain doses of chilling have been provided. The number of days taken for budbreak decreases for buds that have experienced additional chilling. When at least 50 percent of the buds have expanded within twenty-one days of being put into warm conditions, the buds are considered to have fulfilled dormancy requirements, and have now moved into a different phase in the dormancy cycle. Dormancy can be divided into three phases. Paradormancy occurs during the growing season due to factors that arise outside the bud tissue, typically from plant hormone concentrations. For example, hormones produced in the shoot tip prevent lateral growth too close to the tip. This phenomenon, known as apical dominance, dictates the general conical shape of spruce trees (Picea), for instance, because the uppermost buds are more suppressed than the lower buds. Because grapevines are pruned, paradomancy is minimized, allowing lateral branching throughout the growing season. Endodormancy occurs when factors within the bud prevent growth. This phenomenon is triggered by decreasing day length and temperatures in the autumn. Endodormant buds resist growth, even when taken from the outside and placed in a warmer environment. Ecodormancy, as the prefix eco hints, occurs when environmental factors prevent the resumption of growth. Buds transition from endo- to ecodormancy through chill accumulation. Essentially, plants clock how long the winter has been, and this tells them whether they should grow when exposed to warm temperatures (spring has arrived), or if they should remain dormant (midwinter warm spell). Once the buds are ecodormant, they will only remain in a dormant state so long as temperatures remain low. 14 Arnoldia 76\/2 ? November 2018 The authors collect dormant grapevine samples and expose the buds to incremental durations of cold in the laboratory, testing the mechanisms by which plants know to produce new growth (including flower buds, shown above) in the spring. The necessity for chilling is one of the major factors that determines the distribution of vineyards in subtropical and temperate climates. In regions where insufficient chilling occurs naturally, grapevines and other fruit tree species require extra help to transition from endo- to ecodormancy. Wine growers can apply artificial stressors, such as agrochemicals (e.g. hydrogen cyanide), natural compounds (e.g. garlic extract), or even heat from mobile ovens, to jolt endodormant buds into an unnatural ecodormant state, much like a blaring alarm clock. Yet even though these methods can bypass chilling requirements, none are capable of stimulating budbreak as synchronously as natural processes. In addition to dormancy, plant tissues must have mechanisms to cope with below-freezing temperatures. Leaves and other green tissues are generally very sensitive to freezing, due in part to their high water content. This is why deciduous plants lose their leaves in the fall. In the case of grapevines, buds survive the winter through a process called supercooling. The precise mechanisms that contribute to differences in supercooling ability are exceptionally complex and not well characterized, but at the most basic level, supercooling allows water to be cooled below 32?F (0?C) and remain liquid. In fact, given the right conditions, water can remain liquid to temperatures around -40?F (-40?C), although once frozen, the water will only melt at 32?F (0?C). You might have observed this phenomenon if you have ever left a water bottle in your car overnight when the temperature dropped below freezing. When you grab the bottle the following morning, the water may still be liquid, but when you shake the bottle, ice will immediately start forming. Supercooling does not necessitate altering the concentration of sugars or other metabolites--the antifreeze method used in a car engine--rather, with grapevines and many temperate fruit species, physical barriers to ice formation seem to play major roles in the supercooling process. Supercooling provides variable levels of cold hardiness for grapevine buds throughout the winter. If the outside temperature drops below the current level of cold hardiness, ice will form inside the bud tissues, killing the tiny leaves and flowers beneath the bud scales. This means that the threshold temperature for ice formation changes throughout late fall, winter, and early spring. As temperatures begin to dip below freezing in the late fall, grapevine buds slowly Grape Dormancy 15 begin to gain cold hardiness, gradually increasing their ability to survive freeze events. The buds always must remain ahead of the environmental temperature, which is very important, and in New York, the first freeze event of the season rarely occurs before grapevines have gone dormant due to this process of acclimation. Under artificial conditions in growth chambers, we have found that if temperatures are held or oscillated around 27?F (-3?C), dormant buds can acclimate to survive temperatures as low as -4?F (-20?C). But the process isn't instantaneous and may take up to two weeks. Dormancy Danger Understanding the mechanics of dormancy matters because winter is dangerous for grapevines and other perennial plants, and when it comes to agricultural production, predictable harvests are paramount. Depending on the climate of different grape-growing regions, the most perilous time of winter can differ. In New York, slow temperature decline during the fall helps grapevines fully prepare for winter. In contrast, in the Pacific Northwest--an increasingly respected wine region, known for producing European-inspired vintages--the coldest days of the year are often in early December, and rapid temperature changes can zap buds before the acclimation process is finished. Wine growers in New York don't escape unscathed; instead problems arise due to midwinter cold temperatures. Because the vast majority of commercial grape cultivars have been selected from Vitis vinifera, the only grapevine species native to the Mediterranean and therefore adapted for hot summers and mild winters, these cultivars can only survive to a maximum cold temperature of around -4 to -13?F (-20 to -25?C). This temperature range is not unheard of in upstate New York, often causing at least partial bud dieback each winter--the reason vineyards in this region are mostly located around the Finger Lakes and on the shores of the Great Lakes. About sixty wild grape species can be found across North America and Asia, and most have greater maximum cold hardiness than the common grapevine. For example, the fox grape (V. labrusca)--the North American species from which `Concord' grapes were selected--can endure temperatures around -22 to -31?F (-30 to -35?C), or even lower. Amur grape (V. amurensis), which has a broad distribution throughout eastern Asia, may be capable The rates of cold acclimation and deacclimation differ from species to species, and even between different genotypes of the same species. Cold hardiness for cultivars of riverbank grape (Vitis riparia `Bougher') and a common grapevine (V. vinifera `Cabernet Sauvignon') are shown relative to temperature fluctuations in Geneva, New York, throughout months spanning 2017 and 2018. Notice how the riverbank grape--adapted for a colder climate in North America-- prepares more rapidly for more severe winter temperatures. 16 Arnoldia 76\/2 ? November 2018 of surviving even lower levels. But while these species seem to promise natural cold hardiness for breeding efforts, our ability to tap into this genetic variation is relatively limited, given that pure water can only supercool to about -40?F (-40?C). Thus, winemakers in regions where winter temperatures drop below this level must be satisfied with importing grapes. If grapevine buds survive acclimation and midwinter temperatures, the next major event occurs when the buds start losing their cold hardiness as spring temperatures warm. We call this process deacclimation. This is another time of great risk, and climate change promises to make this transition even more unpredictable. Climate models suggest that polar vortex events will become more common in late winter and early spring, catching buds during deacclimation and resulting in lethal damage. Luckily, different species deacclimate at different speeds. The riverbank grape (Vitis riparia) tends to be much faster than the common grapevine (V. vinifera), for instance, while the southern wild grapevine (V. cinerea) seems to be much slower. This difference in northern and southern species probably indicates natural adaptations as a result of their respective winter climates. As a northern species, V. riparia probably evolved a fast deacclimation rate to take advantage of the shorter summers, while V. vinifera and V. cinerea, each more adapted to milder winters and hotter summers, lack the adaptive need to race toward growth. In this way, wild grapevines provide us with the natural adaptive differences needed to learn about cold hardiness and dormancy, and also with the breeding material needed to meet the challenges of a changing climate. Climate change has already reduced the amount of winter chill accumulation in most traditional wine regions. Bordeaux, the largest winegrowing area in France, now receives about 75 percent of the chilling it had in the mid1970s. Our latest research has demonstrated that the speed of early spring development is dependent on chill accumulation, and that we can measure this speed based on deacclimation. So to understand the implications of current climate trends in regions like Bordeaux, more detail must be added to our analogy of dormancy as a night of sleep. When you sleep only one to two hours and get up, it feels like you have not rested at all. Once you hit three hours, every additional hour of sleep feels like a great improvement--think about the difference between four and five hours--although after seven hours, each additional hour provides less energy improvement. We were able to measure a similar phenomenon with the grapevine buds. With minimal chill accumulation, the buds lost their hardiness very slowly, but once they accumulated about eight hundred hours, there was a sharp increase in how responsive they were to warm temperatures. After about twelve hundred hours, however, there was little improvement to responsiveness--the equivalent of surpassing seven hours of sleep. In this sense, the transition between endo- and ecodormacy is gradual, not a clear dichotomy between states. What does this mean for viticulture? Despite the reduced chill accumulation in Bordeaux, mentioned above, the region still receives plenty of chilling for Vitis vinifera cultivars. As a result, the buds and vines are usually ready to begin growing as soon as spring temperatures warm. In 2017, however, unseasonal warmth in April caused early budbreak as plants were very responsive due to high chill accumulation. A subsequent frost occurred in late April and caused extensive damage to vines, reducing the crop by 40 percent compared to the previous year, resulting in one of the lowest yields in the past thirty years. Thus, the future for wine production is complicated from the standpoint of dormancy, cold hardiness, and sustainable viticulture. Climate models predict less chill in warm regions in the future, leading to a need for different cultivars or the use agrochemicals to force vines. In cooler regions, increased chilling temperatures between 32 and 50?F (0 and 10?C) may lead to earlier budbreak for current cultivars, which could be especially detrimental given the increasingly erratic patterns of late winter cold events. Production of wine is not the main concern with a changing climate, given that food production and broader ecosystem stability are at risk. While our research has focused on grapevines, we expect that similar behavior would be seen with many other horticultural and forest MICHAEL DOSMANN Grape Dormancy 17 Gloryvine grape (Vitis coignetiae, accession 63-92*A) at the Arnold Arboretum was wild collected outside of Sapporo, Japan, where record winter lows have hit -19.3?F (-28.5?C) and where average winter snowfall tops seventeen feet (nearly six meters). species. Plant distribution is governed by temperature, and these are generally predicted to increase in the future. This means the optimal zones for many plants will move towards the poles--especially if they require chilling. Agricultural production can more readily adapt with new cultivars and species, but forests may need a hand if we want to continue seeing the current diversity available. Minimum temperatures are the most limiting factor for plant distribution, so as the world gets warmer, it is perhaps a little ironic that increasing our understanding of how plants respond to cold may be key to predicting how they will survive in the future. Additional reading list Cook, B. I., and Wolkovich, E. M. 2016. Climate change decouples drought from early wine grape harvests in France. Nature Climate Change, 6(7): 715?720. Gray, L. K., and Hamann, A. 2013. Tracking suitable habitat for tree populations under climate change in western North America. Climatic Change, 117(1-2): 289?303. Gu, L., Hanson, P. J., Post, W. M., Kaiser, D. P., Yang, B., Nemani, R., and Meyers, T. 2008. The 2007 eastern US spring freeze: Increased cold damage in a warming world?. AIBS Bulletin, 58(3): 253?262. Hannah, L., Roehrdanz, P. R., Ikegami, M., Shepard, A. V., Shaw, M. R., Tabor, G., and Hijmans, R. J. 2013. Climate change, wine, and conservation. Proceedings of the National Academy of Sciences, 110(17): 6907?6912. Kovaleski, A. P., Reisch, B. I., and Londo J. P. 2018. Deacclimation kinetics as a quantitative phenotype for delineating the dormancy transition and thermal efficiency for budbreak in Vitis species. AoB PLANTS, 10(5): ply066. Londo, J. P., and Kovaleski A. P. 2017. Characterization of wild North American grapevine cold hardiness using differential thermal analysis. American Journal of Enology and Viticulture, 68: 203?212. Al Kovaleski is a postdoctoral researcher at the United States Department of Agriculture's research station in Geneva, New York. He completed his doctorate at Cornell University in 2018. Jason Londo is a research geneticist with the United States Department of Agriculture's Grape Genetics Unit, based at the New York State Agricultural Experiment Station in Geneva, New York. He is an adjunct associate professor at Cornell University's School of Integrative Plant Science. "},{"has_event_date":0,"type":"arnoldia","title":"Wood Under the Microscope","article_sequence":3,"start_page":18,"end_page":23,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25654","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060b728.jpg","volume":76,"issue_number":2,"year":2018,"series":null,"season":null,"authors":"Kasia Zieminska","article_content":"18 Arnoldia 76\/2 ? November 2018 Wood Anatomy 19 WOOD U N DE R TH E M IC RO S CO PE Kasia Ziemiska The photo below shows the microscopic structure of twig wood from an American beech (Fagus grandifolia, accession 6832008*A) growing at the Arnold Arboretum. The cross-section is about 10 ?m thick, photographed at 200? magnification. Under the microscope, wood cross-sections have very little contrast, unless stained, and it is difficult to discern some tissues. Here, the lower-right section has been colored to indicate the three main interlocked tissues: fibers, parenchyma, and vessels. F P axial P ray W ood--also called xylem--provides mechanical support for plants, transports and stores nutrients, and transports water from roots to leaves. The proportion, structure, and spatial connectivity of wood tissues varies tremendously across species worldwide, but the functional meaning of much of that diversity remains unclear, leaving much for researchers to discover. FIBERS look like small donuts within a cross- section. But in fact, they are elongated spindle-like cells. Across species worldwide, fibers are the most abundant wood tissue on average, but their proportion ranges from around 10 to 95 percent. In this American beech, fibers occupied 32 percent of twig wood volume and had medium-thick cell walls. Fibers primarily provide mechanical stability for plants--the strength to support limbs and resistance to environmental stresses like wind or snow. PARENCHYMA is a three-dimensional network of living cells, with ray parenchyma connecting bark with inner portions of the stem and axial parenchyma running parallel with the length of the stem. This network stores and transports nutrients, which provide energy for building and maintaining living tissues. Parenchyma also contributes to pathogen defense, and it may factor into water storage and the continuity of water transport, although the exact mechanisms of these functions are yet uncertain. Parenchyma occupies between 5 and 90 percent of wood volume for species globally. In this American beech, parenchyma occupied 30 percent. VESSELS are like pipes that transport water from roots to leaves. Their diameter is larger than other wood cells, and they differ tremendously across species, ranging from around 15 to 250 ?m. The diameter has a huge influence on how efficiently water can be transported through wood. The larger the vessel, the more efficient the transport. In this American beech, average vessel diameter was 20 ?m. V 100 m F fibers P parenchyma V vessels 20 Arnoldia 76\/2 ? November 2018 Wood samples from seven tree species at the Arnold Arboretum illustrate striking differences in wood anatomy, even among trees adapted for similar climates. Notice the small but relatively consistent diameter of vessels in Liriodendron, compared to the prominent seasonal variation in the Quercus sample. Likewise, other tissues show similar diversity of dimension and distribution. Photos are all at the same scale, so cell sizes can be compared. Each image shows one or more growth rings. fibers F parenchyma P vessels V growth ring boundary G G G P F G V Osage orange (Maclura pomifera, 79-46*D) G G V G F F V P P G 100 m Tulip tree (Liriodendron tulipifera, 14992*A) G V Northern catalpa (Catalpa speciosa, 927-58*B) Wood Anatomy 21 V P G V P F F V V G Honeylocust (Gleditsia triacanthos, 14681*A) G V Chinkapin oak (Quercus muehlenbergii, 389-91*C) G P V F P V V G Korean paulownia (Paulownia tomentosa `Coreana', 730-77*D) F G Common persimmon (Diospyros virginiana, 14513*B) 22 Arnoldia 76\/2 ? November 2018 THE ANATOMY OF TREE LIFE T here are myriad ways plants can make a living. Herbaceous annuals grow fast and live short, while other plants, like trees, grow slow and live long. Plants can produce cheaply built leaves that catch sunshine for only a few months, and others produce robust leaves that survive for several years. In a forest, shade-tolerant trees produce short stems, while sun-thirsty ones stretch tall. This marvelous diversity of form and function persists at the microscopic level, hidden to the naked eye yet biologically consequential. Wood anatomy is not a new discipline. It has been studied since the seventeenth century, when Robert Hooke famously turned a microscope to a sample from the cork oak (Quercus suber) and coined the term cell based on the walled structures he observed. Shortly thereafter, in 1673, Antonie van Leeuwenhoek began observing the microscopic structure of cross-sectioned twigs from numerous woody species, including temperate and tropical plants. Over the next several decades, he rendered remarkably detailed illustrations, which are among the earliest progenitors to the images shown on the following pages. Yet the intensity and emphasis of these anatomical studies has changed with time. An increased interest in wood anatomy in the nineteenth century was related to taxonomy-- classifying differences between groups of plants--and Harvard University and other institutions built large collections of wood samples (usually rectangular blocks), which allowed researchers to easily compare wood from various species. This descriptive research continued into the twentieth century, although most of the studies were of qualitative nature, where anatomical structures were described using categories like \"thin\" and \"thick,\" instead of unit measurements. Over the past sixty years (and especially over the last two decades), however, an increased number of researchers started quantifying anatomical features, and importantly, they began considering the relationship between structure and function of wood on an ecological and climactic scale--attempting to understand why species adapted to disparate habitats possess different wood anatomies. The three main wood tissues--fibers, parenchyma, and vessels--are also present in herbaceous plants, simply in different organizations. These tissues play several vital functions. They transport water and nutrients, while also providing nutrient storage, pathogen defense, and mechanical support. We are still learning about additional roles; for example, scientists have found that nutrients stored in wood might be used for maintenance of continuous water transport, not only for building new organs like bark and leaves. Of the three main tissues, researchers have the greatest understanding of vessels, the cells responsible for transporting water. The average size of vessels varies hugely worldwide, from around 15 to 250 ?m in diameter. We know that large vessels tend to be more susceptible to embolism--air blockage in a vessel--which impedes water transport and, if common, can be lethal. Embolism can be caused by freeze-thaw events or drought, so species exposed to such stresses tend to have smaller vessels. On the other hand, smaller vessels are less efficient for water transport. Beyond vessel diameter, plants can make many other anatomical adjustments to alter their hydraulic functioning. Some plants have lots of vessels, others very few, and plants can change the size of these structures across the growing season. (On the previous pages, large vessels can often be observed at the beginning of each growing season, when plenty of water was abundant, and the dimensions progressively shrink throughout Wood Anatomy 23 the season.) Plants can also change how vessels are interconnected via the shape and size of small openings called pits. Because anatomical measurements are time consuming, studies that look at highly detailed anatomy are usually limited to, at most, a few dozen species. Recent use of metaanalysis, however, has combined data from many individual studies, shedding light on anatomical variation worldwide--primarily for vessel diameter, vessel proportion, and parenchyma proportion. As a Putnam Fellow at the Arnold Arboretum, my research focused on parenchyma, the tissue that is understood to store and transport nutrients, along with providing pathogen defense and water storage. One of the large-scale analyses had recently revealed that, across more than fifteen hundred species worldwide, the proportion of parenchyma in wood varies remarkably, ranging from 5 to 90 percent of the total volume (Morris et al., 2016). Species with the highest parenchyma proportions are only found in the tropics, yet species with little parenchyma are found in all climates: temperate, subtropical, and tropical. The functional meaning of this geographical distribution remains unclear, however, which is astonishing considering the dramatic extent of the variation. Given the traditional understanding that parenchyma stores water, I investigated the hypothesis that woody species with a higher proportion of parenchyma would have greater water-storage capacity, allowing them to withstand drought more readily--essentially like having more water bottles in the pantry. Across thirty tree species that I studied in the Arnold Arboretum, however, species with more parenchyma didn't necessarily store additional water, nor were they better at accessing whatever water had been stored. So the mystery of parenchyma proportion in wood continues to be unresolved. Presumably species with little parenchyma have entirely different life strategies than species with lots of parenchyma. This information will be important for helping us predict how plants will respond to increased aridity in many areas under climate change. An anatomical perspective within plant sciences is essential for unravelling ecological and physiological questions about how species evolved to grow where they do. William Louis Stern--a professor at the University of Maryland--noted the need for this interdisciplinary approach in the pages of Arnoldia back in 1973. \"Lest I am accused of being a wood anatomy bigot,\" he wrote, \"let me hasten to say that I do not believe that studies in wood anatomy can remain viable in a vacuum; rather, they must be integrated with other studies in plant anatomy and with other phases of botanical endeavor.\" Wood anatomy is timeconsuming and slow--requiring patience and meticulous microscopy skills--but the more we know, the more we realize that we still have much to learn. Work Cited Morris, H., L. Plavcov?, P. Cvecko, E. Fichtler, M.A. Gillingham, H.I. Mart?nez-Cabrera, D.J. McGlinn, E. Wheeler, J. Zheng, K. Ziemiska, and S. Jansen. 2016. A global analysis of parenchyma tissue fractions in secondary xylem of seed plants. New Phytologist, 209: 1553?1565. Kasia Ziemiska is an Arnold Arboretum research associate and a recent postdoctoral Putnam Fellow. "},{"has_event_date":0,"type":"arnoldia","title":"Old Molecules, New Climate: <i>Metasequoia</i>'s Secrets","article_sequence":4,"start_page":24,"end_page":32,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25652","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060af6b.jpg","volume":76,"issue_number":2,"year":2018,"series":null,"season":null,"authors":"Leng, Qin; Yang, Hong","article_content":"Old Molecules, New Climate: Metasequoia's Secrets Hong Yang and Qin Leng A round forty-five million years ago, the Arctic was ice free, scarcely the expanse of lichen-encrusted rock and glaciers that we see there today. Fossil records reveal that an extensive forest flourished throughout the early Cenozoic, when the canopy was predominated by Metasequoia and other deciduous conifers. The single remaining species of this genus, M. glyptostroboides, is known as the dawn redwood and is now restricted to a small population in south-central China, around forty-two hundred miles (sixty-eight hundred kilometers) south of this historic distribution. When botanists first learned about the living population more than seventy years ago, no one could have imagined that those plants would provide crucial clues for understanding more than one hundred million years of historic climate change--not to mention changes to come. Yet the rare discovery of fossils containing exquisitely preserved organic tissues and biomolecules, coupled with new molecular research techniques, has revealed just that. Traditional paleobotanical studies are comparative--drawing links between the anatomy of fossils and their living relatives--while molecular analyses of isotopes and biomolecules (such as lipids, carbohydrates, and lignin) are usually reserved for modern samples of freshly harvested material. Recent innovations with laboratory instruments, however, have made it possible for researchers, including ourselves, to extract valuable molecular information from so-called rocks. With Metasequoia, which boasts a long fossil record, the implications of this research are especially pronounced given that we can test hypotheses at the molecular level across an enormous timescale. The Green Arctic Scientists (and museumgoers) usually encounter two types of plant fossils: either imprints or compressions. Imprints are analogous to animal tracks, occurring when plant tissue remains pressed into sediments and subse- quently decomposes, while compressions occur when the tissue becomes sandwiched between flattened layers of sedimentary rocks. In the late 1990s, however, Ben LePage and Chris Williams, then working at the University of Pennsylvania, showed us three-dimensionally preserved Metasequoia fossils that they had collected from the Canadian islands of Ellesmere and Axel Heiberg, on the northwestern side of Greenland. We could hardly believe that the fossils were from Cenozoic strata that was around fifty millions years old. The dark-brown leaves, stems, and cones resembled a pile that someone had raked in their backyard--loose but delicate, soft but brittle, individually separated but tightly packed. When Metasequoia lived in the warm and humid Arctic, the plants shed their leaves in autumn, before the four months of total darkness. The litter was buried for tens of millions of years, including beneath the weight of a continental ice cap. Ironically, the tundra surface in the high Arctic has been increasingly exposed due to recent global warming, revealing some of the best fossil material to study ancient climate change. The unrivaled quality of these mummified Metasequoia fossils extends beyond what was visible to the naked eye. Using molecular technologies, we detected biomolecules--plentifully! Although cellulose and other polysaccharides are abundant in living plant tissues, they are rarely found in ordinary fossils. Preserved for around fifty million years, these Metasequoia fossils possess the oldest biomolecules of this kind ever recovered. Moreover, our molecular evidence suggests that these molecules actually provide both the physical enforcement and chemical stability for maintaining the three-dimensional structures of these exceptional fossils. The Arctic plays a crucial role in the dynamics of Earth climate. Not only does it actively influence climate on a global scale, but the Arctic passively receives climate change feedback as well. Interestingly, Metasequoia has been a witness CHRISTOPHER J. WILLIAMS QIN LENG AND HONG YANG CHRISTOPHER J. WILLIAMS Metasequoia 25 Fossilized Metasequoia trunks from the late Paleocene and early Eocene have been discovered on Ellesmere Island, Canada, along with three-dimensionally preserved leafy fossils, from which the authors have extracted biomolecules and isotope signals. The leafy branchlet in the upper right corner measures 0.83 inches (2.1 centimeters) long. 26 Arnoldia 76\/2 ? November 2018 and faithful recorder of such changes. At Axel Heiberg and Ellesmere Islands, with latitudes higher than 80? north, large amounts of fossilized Metasequoia trunks--up to twenty-six feet (eight meters) long and ten feet (three meters) in diameter at breast height--indicate the large size of these forests, which were nurtured by a warm and humid high-latitudinal climate. Williams and colleagues demonstrated that the productivity of these dense Metasequoia forests was comparable with modern-day temperate rainforests, like the rich landscape of the Olympic Peninsula in Washington State. Yet the Metasequoia landscape was primarily populated with deciduous plants, not evergreens--evidence of adaptions for four months of complete darkness at the northern latitudes. Animal fossils are rarely found with these Metasequoia remains, but fossils found in similar-aged strata at Ellesmere Island include rhinoceros- and hippo-like mammals, along with giant tortoises and alligators--all indicative of a humid swampy environment, consistent with the reconstruction by plant fossils. Clearly, the ice-free Arctic during the early Cenozoic was a completely different world compared with the barren landscape today. Living Climate Legacies To understand the climate that supported such impressive forests in the Arctic, however, our ability to study a living Metasequoia species is essential. The first climate-related experiment involving the living species, M. glyptostroboides, was launched inadvertently. When trees were discovered in western Hubei Province, China, in 1944, two American scientists immediately recognized the importance of this plant: Ralph Chaney from the University of California, Berkeley, and Elmer Merrill, the director of the Arnold Arboretum. Through separate collaborations with Chinese colleagues-- notably Wan-Chun Cheng and Hsen-Hsu Hu-- both Chaney and Merrill arranged for the collection of dawn redwood seeds from China and distributed them to botanical gardens around the United States and Europe. Despite their arguments about who should get the credit for making these seeds available for cultivation, the resulting widespread dispersal to gardens in dra- matically different climatic zones (essentially all corners of the contiguous United States) set up a natural experiment that we have dubbed the Chaney-Merrill Experiment. The remnant population of Metasequoia in China represents a relatively homogenous genotype, so we can obtain information about how the seedlings survived in disparate gardens, and eliminate some confusion about nature versus nurture. Physiological studies based upon samples from these trees revealed that M. glyptostroboides can endure a wide range of climate conditions. We have obtained leaf samples from forty trees across this range, and these are stored in our Laboratory for Terrestrial Environments at Bryant University, tightly packed in yellow envelopes and frozen. These leaf tissues have helped us generate systematic molecular and biochemical data, which we compared with climate data from the past seventy years from the locations where the trees have been growing. We found that biomolecular compositions within the leaves changed relative to latitude, average temperatures, and average annual precipitation. These correlations established a necessary baseline for interpreting biomolecules and biochemicals that we would later obtain from Metasequoia fossils. Another recent experiment on Metasequoia glyptostroboides provided additional context. Richard Jagels and his colleagues and students at the University of Maine designed a greenhouse experiment to examine how the genus would have performed within its historic distribution inside the Arctic Circle. The climate in this northern region was temperate during the early Cenozoic, yet the unique light regime would have remained consistent: up to four months of complete darkness and four months of twenty-four-hour sunshine. To test the physiological adaptation of M. glyptostroboides--as a living stand-in for the Cenozoic species, M. occidentalis--to this light regime, the team partitioned a large greenhouse on their campus into two different compartments: one with normal light, corresponding to the middle latitude of Maine (45? north), and the other with continuous light, mimicking the same low angle and low intensity of Arctic light conditions during summer months. MICHAEL DOSMANN KYLE PORT DANNY SCHISSLER Metasequoia 27 Metasequoia glyptostroboides is the only extant member of a genus that once flourished at warm Arctic latitudes during the early Cenozoic. Clockwise from top left: Seed cones hang on winter branches (notice that the deciduous leaves have already fallen), small pollen cones emerge in the spring, and trees at the Arnold Arboretum showcase an affinity for moist habitats. ARNOLD ARBORETUM AND GIS COMMUNITY 28 Arnoldia 76\/2 ? November 2018 The Canadian islands of Ellesmere and Axel Heiberg are among the most northerly landmasses on the planet, occurring well above the Arctic Circle. Metasequoia fossils have been recovered from both islands, at latitudes above 80?N. Seedlings of Metasequoia glyptostroboides were grown along with those of two other deciduous conifers: bald cypress (Taxodium distichum) and tamarack (Larix laricina), whose fossil relatives were also common in warm Arctic floras during the early Cenozoic. These plants were grown for two consecutive years under the two different light regimes with otherwise identical conditions, including temperature, relative humidity, carbon dioxide level, and greenhouse irrigation. This research revealed that, in addition to deciduous leaves, which would drop during the prolonged darkness of Arctic winters, Metasequoia possesses physiological characteristics, such as high photosynthesis capacity and improved water-use efficiency, that help it take advantage of the weak but continuous Arctic summer light. This helps explain how this genus outcompeted other plants in the warm Arctic. At the end of the two-year Jagels Experiment, we corresponded with the researchers and obtained leaf tissue from Metasequoia seedlings grown in these greenhouse conditions. We wanted to learn how the ratios of carbon and hydrogen isotopes--slight variations of these elements built into plant tissues through photosynthesis--changed under different light treatments. These isotopes have been commonly used to understand ancient patterns for temperature, precipitation, and carbon dioxide level, and indeed, we discovered noticeable differences between seedlings grown under these alternate light regimes. Even more importantly, our work, published in 2009, established precise empirical relationships between isotope values of plant lipids and environmental water, allowing us to infer ancient moisture levels in this Arctic habitat. New Technologies for Old Molecules Experiments based on fresh Metasequoia glyptostroboides samples have enhanced the ability of researchers to interpret data from M. occi- Metasequoia 29 dentalis fossils, including biomolecules stored within the seasonal growth rings of the large trunks on Ellesmere and Axel Heiberg Islands. Hope Jahren--a geochemist at the University of Oslo, now well-known for her science memoir, Lab Girl--conducted analyses on seasonal variation of isotopes within these rings. Jahren and her colleague Leonel Sternberg observed highresolution patterns in these isotopes, allowing them to reconstruct the impact of subtle climate variations on the growth habit of Metasequoia in the warm Arctic. They estimated a mean annual temperature for this high-latitude region to be around 55?F (13?C) during the Eocene-- about double the present-day measurements. Relative humidity estimates were equally high: around 67 percent during the growing season and close to 100 percent towards the end of the growing season. This climate information has direct implications for understanding global precipitation patterns during the early Cenozoic. At present, Arctic ice reflects back large amounts of solar radiation, keeping the global temperature low and simultaneously creating a steep temperature gradient across different latitudes. This equator-to-pole temperature difference significantly impacts the general circulation of heat--and moisture--through the atmosphere and ocean. Given what we know about Arctic temperatures, relative humidities, and carbon dioxide levels during the early Cenozoic, we know that this gradient of temperatures between latitudes would have been less pronounced, significantly impacting precipitation patterns. We used the relationship between environmental water and isotopes in fossilized Metasequoia leaves (established with the Jagels Experiment) to propose a model for early Cenozoic moisture patterns in the Arctic. To our surprise, the relatively low hydrogen isotope values we measured were not compatible with the conventional understanding that the reduced temperature gradient from the equator to the Arctic should result in less precipitation during the long-distance transport of moisture within the atmosphere, depositing water with heavier hydrogen within the Arctic. Although there is no modern analogue for these high-latitude forests, dense forests at lower latitudes, such as the temperate rainforests, offer clues that could explain these low isotope measurements. Due to moisture generated through extensive evapotranspiration of the vegetation, a portion of the heavy precipitation above these dense forests is composed of locally recycled moisture with lighter hydrogen. The greenhouse simulation in the Jagels Experiment supported this interpretation, demonstrating that photosynthesis under four-months of continuous light enhanced water evapotranspiration. These observations suggest that Metasequoia forests had a dynamic impact on moisture patterns in the ancient Arctic and may, in that sense, have even played an important role in maintaining the air circulation at the ice-free Arctic. Climate Predictions Strikingly, it took millions of years for the Arctic to transform from a humid Metasequoia-dominated forest into the landscape we recognize today, but the inverse warming trend now appears to be happening at a much faster rate. Over the last thirty years, Arctic warmth has accelerated along with rising carbon dioxide levels in the atmosphere. Arctic sea ice is melting, and glaciers are retreating at an unprecedented pace. In 2017, sea ice reached the lowest extent since the earliest time of satellite measurement in the 1960s. Recent global circulation modeling suggests that if the warming trend continues, by as early as the 2030s, the Arctic Ocean will change from perennially ice-covered to seasonally ice-free, further decreasing the temperature gradient between the equator and the poles. The weakened moisture delivery towards the poles will likely alter storm trajectories and increase temperature and precipitation anomalies, affecting the life of plants and animals, humans included, at the global scale. In addition to informing what we know about ancient climate, Metasequoia has also contributed to future climate models. The scientific community has long accepted that atmospheric carbon dioxide has been one of the primary drivers for global temperature changes, but the rate at which temperature increases in response to changes of this greenhouse gas--a metric known YUYANG ZHUGE 30 Arnoldia 76\/2 ? November 2018 An artistic reconstruction of a Metasequoia-dominated forest and its surrounding environment at the end of the Arctic growing season during the middle Eocene (around 45 million years ago). The composition and density of the forest, as well as the hydrological conditions, were based upon fossil records and their stable isotope analyses. as climate sensitivity--has been the subject of significant and ongoing research. Paleoclimatologists do not have the luxury of directly measuring ancient carbon dioxide levels; instead, they rely on indirect estimates, known as proxies, to make an inference. One of the most reliable proxies for reconstructing atmospheric carbon dioxide levels for geological eras that predate the oldest ice-core records is stomatal frequency. Stomata are small openings on the surface of leaves, and are the means through which terrestrial plants control the balance between absorbing carbon dioxide and losing water into the air. Species-specific relationships between stomatal frequency and atmospheric carbon dioxide (under high carbon dioxide levels, plants produce fewer stomata) thus allow scientists to predict one from the other. The ideal plant to study this phenomenon should have a continuous, abundant, and widespread fossil record, along with living representatives to provide detailed comparative analyses and calculation. The genus Metasequoia fits the bill perfectly. The only challenge is that, as a deciduous conifer, Metasequoia has a very thin and fragile cuticle--the waxy layer covering its leaves--making it difficult to calculate stomatal frequency from fossils. Recent studies using improved experimental treatments and bioimaging techniques of Metasequoia fossils were successful, however, and surprisingly, work by Daniel Maxbauer and colleagues, based upon Metasequoia fossils from the Axel Heiberg Island and published in 2014, found that carbon dioxide levels during the middle Eocene (about thirty-seven to forty-seven million years Metasequoia 31 Metasequoia's Legacy Continues As we look toward future research involving Metasequoia, we can't help but marvel over the unintended consequences of scientific discoveries. When Japanese paleobotanist Shigeru Miki made the seemingly routine description of a new fossil species belonging to an extinct genus in 1941, he had no idea that its living equivalent was growing a few thousand miles away and waiting to be discovered. When Chinese botanist Zhan Wang encountered a splendid dawn redwood tree in Hubei Province (then part of Sichuan) for the first time, he could not have predicted that this rare conifer would produce important clues for understanding the vexing problem of climate change. From the Chaney-Merrill Experiment to the Jagels Experiment, from the discovery of exceptional fossils in the Arctic to the applications of molecular isotope technology, science, as illustrated with Metasequoia, is a continuous endeavor in which new technologies facilitate new questions and, ultimately, new breakthroughs. Whether inconspicuously planted along a roadside in Sydney, Australia, or proudly JONATHAN FONSECA ago) may have been about half of what were previously estimated using other methods: a median of 424 parts per million (ppm) for the middle Eocene. If accurate, this work suggests that the Metasequoia forests in the Arctic were supported by much lower atmospheric carbon dioxide levels than previously thought. This scenario does not portend good news for humans. The latest data from the National Oceanic and Atmospheric Administration show that carbon dioxide levels reached 405 ppm in 2017, exceeding concentrations throughout the past eight hundred thousand years that can be observed using ice cores. At the time when this article was written (in the summer of 2018), the level reached 410 ppm, marking the highest level since the late Pliocene, around three million years ago. From the pre-industrial carbon dioxide levels between 260 and 270 ppm to the current level, it took us less than two centuries to increase the atmospheric carbon dioxide level by more than 100 ppm. With this alarmingly accelerated rate, it wouldn't take long to reach levels that once sustained dense deciduous forests across an ice-free Arctic. The authors examine Metasequoia glyptostroboides on the Bryant University campus. These trees were planted in 2006 during the Second International Metasequoia Conference held at Bryant and Yale. 32 Arnoldia 76\/2 ? November 2018 showcased on our campus at Bryant University in Rhode Island, the dawn redwood has thrived through cultivation around the globe. In terms of sheer numbers, Metasequoia seems to have survived from the brink of extinction, yet its native population remains isolated with low levels of genetic diversity. While the natural population's long-term survival remains uncertain under the changing climate, what is certain is that, along with the advancement of technology, both living and fossil Metasequoia will continue to offer us invaluable information about its past secrets and the future of our global climate. Acknowledgement With limited pages here, we can only highlight the many exciting scientific inquiries about Metasequoia contributed over the past two decades. This is the duration since Hong Yang's first Arnoldia article on Metasequoia fossils and molecules, which was published in a 1998 special issue celebrating the genus. Readers can obtain further reading from the references and especially the proceedings of the three International Metasequoia Conferences. We would like to thank Jonathan Damery for discussing the structure and editing the manuscript, Chris Williams and Jonathan Fonseca for providing photograph images, and Yuyang Zhuge for illustrating the Eocene Arctic Metasequoia forest and its surrounding environment based on the scientific data we provided. References Jahren, A.H. 2007. The Arctic forest of the Middle Eocene. Annual Review of Earth and Planetary Sciences, 35: 509?540. Jahren, A.H., and L.S.L. Sternberg. 2008. Annual patterns within tree rings of the Arctic middle Eocene (ca. 45 Ma): Isotopic signatures of precipitation, relative humidity, and deciduousness. Geology, 36: 99?102. Jagels, R., and M.E. Day. 2004. The adaptive physiology of Metasequoia to Eocene high-latitude environment. In: A.R. Hemsley and I. Poole (Editors). The Evolution of Plant Physiology. Elsevier Academic Press, 401?425. Leng, Q., G. Langlois, and H. Yang. 2010. Early Paleogene Arctic terrestrial ecosystems affected by the change of polar hydrology under global warming: Implications for modern climate change at high latitudes. Science China Earth Science, 53: 933?944. LePage, B.A., C.J. Williams, and H. Yang (Eds.) 2005. The geobiology and ecology of Metasequoia. Springer Netherlands, Dordrecht, the Netherlands; Norwell, MA, USA. (The proceedings of the First International Metasequoia Conference) Maxbauer, D.P., D.L. Royer, and B.A. LePage. 2014. High Arctic forests during the middle Eocene supported by moderate levels of atmospheric CO2: Geology, 42(12): 1027?1030. Noshiro, S., Q. Leng, B.A. LePage, A. Momohara, H. Nishida, K. Uemura, C.J. Williams, and H. Yang (Eds.) 2011. Metasequoia: The legacy of Dr. Shigeru Miki. Proceedings of the Third Inter national Metasequoia Symposium. Japanese Journal of Historical Botany, 19: 1?136. Wang, Y.Q., A. Momohara, L. Wang, J. Lebreton-Anberr?e, and Z.K. Zhou. 2015. Evolutionary history of atmospheric CO 2 during the Late Cenozoic from fossilized Metasequoia needles. PLoS ONE, 10(7): e0130941. Williams, C.J., A.H. Johnson, B.A. LePage, D.R. Vann, and T. Sweda. 2003. Reconstruction of Tertiary Metasequoia forests. II. Structure, biomass, and productivity of Eocene floodplain forests in the Canadian Arctic. Paleobiology, 29: 271?292 Yang, H. 1998. From fossils to molecules: the Metasequoia tale continues. Arnoldia, 58(4) and 59(1): 60?71. Yang, H., and L.J. Hickey (Eds.) 2007. Metasequoia: Back from the brink? An update: Proceedings of the Second International Symposium on Metasequoia and Associated Plants. Bulletin of the Peobody Museum of Natural History, 48: 183?426. Yang, H., and Q. Leng. 2009. Molecular hydrogen isotope analysis of living and fossil plants: Metasequoia as an example. Progress in Natural Science, 19: 901?912. Yang, H., M. Pagani, D.E.G. Briggs, M.A. Equiza, R. Jagels, Q. Leng, and B.A. LePage. 2009. Carbon and hydrogen isotope fractionations under continuous light: Implications for palaeoenvironmental interpretations at high Arctic during Paleogene warming. Oecologia, 160: 461?470. The map in this article was created using Esri, USGS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, lntermap and the GIS user community. Hong Yang (hyang@bryant.edu) is Charles J. Smiley Professor of Geobiology, and Qin Leng (qleng@bryant.edu) is Professor of Biology; both co-direct the Laboratory for Terrestrial Environments at Bryant University in Rhode Island, USA, where Metasequoia has been a subject of investigation for various research projects related to both modern and ancient environments. "},{"has_event_date":0,"type":"arnoldia","title":"Overstory View","article_sequence":5,"start_page":33,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25653","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060b36f.jpg","volume":76,"issue_number":2,"year":2018,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"BOOK REVIEW Overstory View Jonathan Damery R ecord tags at the Arnold Arboretum are as much a record of human lives as the plants. The oldest living accession--the sole remnant from 1872, the year the Arboretum was founded--comprises three shrubby winterberry hollies (Ilex verticillata, accession 147*A, C, and E), tucked near the North Woods. These were collected near Boston by Jackson Dawson, the Arboretum's first plant propagator, who was then overseeing greenhouses for the recently established Bussey Institute. If you indulge philosophical predilections, it is awe-inspiring to eye these wane suckering shrubs, which bore a smattering of red fruit this year, and realize that the shrubs have now persisted more than a century beyond the man who collected the original seed. As a rule, the Arboretum's oldest plants have all outlived their collectors, and at least some members of the subsequent plant generations, even material collected on our four expeditions this year, are destined to do the same. Richard Powers explores this premise in his new novel, The Overstory. The narrative structure flexes time and space, at first introducing characters that exist in different decades (and centuries) and in different parts of the United States. All of the characters develop affinities for specific trees. In some cases, the relationship becomes a family legacy, as with the introductory characters: European immigrants named J?rgen and Vi Hoel move from Brooklyn to the Iowa prairie in the mid-nineteenth century. J?rgen plants an American chestnut (Castanea dentata) from seed he brought west. After J?rgen's death, his son buys a Kodak No. 2 Brownie--the classic model once used by hobbyist photographers--and, in 1903, he begins photographing the now prospering chestnut from the same location each month. A grandson continues the tradition, even as chestnut blight (Cryphonectria parasitica) sweeps through the eastern population, and the lens finally passes to a great-great-grandson named Nicholas. When Nicholas looks at the photos his family has produced, \"three-quarters of a century dances by in a five-second flip.\" Moreover, he sees a compression of human existence: \"the holidays of his childhood, the entire clan gathering for turkey or carols, midsummer flags and fireworks.\" This generational project, in the nonfiction realm, is reminiscent of Chinese botanist Kaipu Yin's rephotography of trees and landscapes that Arboretum plant collector Ernest Henry Wilson documented in China in the early twentieth century. The resulting side-by-side comparisons were published in a collection, Tracing One Hundred Years of Change: Illustrating the Environmental Changes in Western China. The passage of a ARNOLD ARBORETUM ARCHIVES 34 Arnoldia 76\/2 ? November 2018 A sixteen-foot slab of this giant sequoia (Sequoiadendron giganteum) was harvested from Kings River Grove, California, in 1891, as part of the Jesup Wood Collection, a project overseen by Charles Sprague Sargent. The slab can still be viewed at the American Museum of Natural History in New York City, where the annual growth rings reveal the tree began growing around 550 AD. Sargent, the founding director of the Arboretum, died in 1927. century is evident in the landscapes and, more specifically, in the form of individual trees. Some have survived and, given their size at the time Wilson photographed them, likely survived at least the century before that. Others, however, have gnarled to a fist of limbs, broken and beaten. Still others are ghosts, replaced with glowing storefronts. This effort to showcase non-human time is even more dramatic with Rachel Sussman's photography collection The Oldest Living Things in the World, because she only includes organisms (mostly plants) that are more than two thousand years old. Of course, one automatically thinks of bristlecone pines (Pinus longaeva), growing as bony skeletons in the White Mountains of California, where some specimens are more than five Book Review 35 thousand years old--a duration that pre-dates the invention of Sumerian script. But Sussman also directs attention to a colony of quaking aspen (Populus tremuloides) in south-central Utah, which has been spreading via underground roots for an estimated eighty thousand years. This marks the approximate time when humans began to successfully migrate from Africa--an astounding timeline, especially considering evolutionary anthropologists have shown that Neanderthals (Homo neanderthalensis) persisted in Eurasia for another forty thousand years beyond this. Incidentally, Powers doesn't miss this ancient colony of aspen. One of his characters, Patricia Westerford, flees academic pariahdom (after publishing a controversial article about plant communication) and drives to the magisterial forests of the Pacific Northwest, but not before stopping to see the Utah aspen. \"The thing is outlandish,\" Powers writes, \"beyond her ability to wrap her head around.\" Likewise, Powers uses narrative to force readers beyond their personal footsteps--to literally view human existence from the overstory of time--which many environmental theorists suggest is essential for conceptualizing issues like climate change, with implications stretching far into the geologic future. Plants at the Arboretum--even lowly shrubs like Dawson's winterberries--provide a legible reminder of organismal time, a scale beyond ourselves, and as Powers demonstrates, narrative has potential to do the same. Books referenced Powers, R. 2018. The Overstory. New York: W. W. Norton and Co. Sussman, R. 2014. The Oldest Living Things in the World. Chicago: University of Chicago Press. Yin, K. 2010. Tracing One Hundred Years of Change: Illustrating the Environmental Changes in Western China. Beijing: Encyclopedia of China Publishing House. Jonathan Damery is the associate editor of Arnoldia. 36673667 U.S. POSTAL SERVICESTATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION(Required by 39 U.S.C. 3685) 1. Publication Title: Arnoldia. 2. Publication No: 0004?2633. 3. Filing Date: October 17, 2018. 4. Issue Frequency: Quarterly. 5. No. of Issues Published Annually: 4. 6. Annual Subscription Price: $20.00 domestic; $25.00 foreign. 7. Complete Mailing Address of Known Office of Publication: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 8. Complete Mailing Address of Headquarters of General Business Office of Publisher: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 9. Full Names and Complete Mailing Address of Publisher, Editor, and Managing Editor: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500, publisher; Jonathan Damery, Arnold Arboretum, 125 Arborway, Boston, MA 02130?3500, associate editor. 10. Owner: The Arnold Arboretum of Harvard University, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 11. Known Bondholders, Mortgagees, and Other Security Holders Owning or Holding 1 Percent or More of Total Amount of Bonds, Mortgages, or Other Securities: none. 12. The purpose, function, and nonprofit status of this organization and the exempt status for federal income tax purposes have not changed during the preceding 12 months. 13. Publication Name: Arnoldia. 14. Issue Date for Circulation Data Below: TBD. 15. Extent and Nature of Circulation. a. Total No. Copies. Average No. Copies Each Issue During Preceding 12 Months: 1,600. Actual No. Copies of Single Issue Published Nearest to Filing Date: 1,600. b. Paid and\/or Requested Circulation. (1) Paid\/Requested Outside-County Mail Subscriptions. Average No. Copies Each Issue During Preceding 12 Months. Copies Each Issue During Preceding 12 Months: 27. No. Copies of Single Issue Published Nearest to Filing Date: 25. (2) Paid In-County Subscriptions. Average No. Copies Each Issue During Preceding 12 Months. Copies Each Issue During Preceding 12 Months: 1,131. No. Copies of Single Issue Published Nearest to Filing Date: 1,109. (3) Sales Through Dealers and Carriers, Street Vendors, and Counter Sales: none. (4) Other Classes Mailed Through the USPS: none. c. Total Paid and\/or Requested Circulation. Average No. Copies Each Issue During Preceding 12 Months: 1,158. Actual No. Copies of Single Issue Published Nearest to Filing Date: 1,044. d. Free Distribution by Mail. Average No. Copies Each Issue During Preceding 12 Months: 141. Actual No. Copies of Single Issue Published Nearest to Filing Date: 140. e. Free Distribution Outside the Mail: Average No. Copies Each Issue During Preceding 12 Months: 250. Actual No. Copies of Single Issue Published Nearest to Filing Date: 250. f. Total Free Distribution: Average No. Copies Each Issue During Preceding 12 Months: 391. Actual No. Copies of Single Issue Published Nearest to Filing Date: 390. g. Total Distribution: Average No. Copies Each Issue During Preceding 12 Months: 1,549. Actual No. Copies of Single Issue Published Nearest to Filing Date: 1,434. h. Copies Not Distributed. Average No. Copies Each Issue During Preceding 12 Months: 51. Actual No. Copies of Single Issue Published Nearest to Filing Date: 166. i. Total. Average No. Copies Each Issue During Preceding 12 Months: 1,600. Actual No. Copies of Single Issue Published Nearest to Filing Date: 1,600. j. Percent Paid and\/or Requested Circulation. Average No. Copies Each Issue During Preceding 12 Months: 75%. Actual No. Copies of Single Issue Published Nearest to Filing Date: 73%. I certify that all information furnished on this form is true and complete. Jonathan Damery, Associate Editor. "},{"has_event_date":0,"type":"arnoldia","title":"Hidden Gem Among Vines: Actinidia arguta","article_sequence":6,"start_page":36,"end_page":37,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25651","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060af26.jpg","volume":76,"issue_number":2,"year":2018,"series":null,"season":null,"authors":"Brinkman, Rachel A.","article_content":"36 Arnoldia 76\/2 ? November 2018 Hidden Gem Among Vines: Actinidia arguta Rachel A. Brinkman T he hardy kiwi (Actinidia arguta) is a vigorous vine with fruits that you are unlikely to find at your local grocery store. The grape-sized nuggets are like the large kiwifruit (A. deliciosa), simply smaller and hairless. When you cut open the dusky fruit, which sometimes blushes from green to red, you'll see a firework design of lime-colored flesh with an inner ring of tiny chocolate-brown seeds. The taste of A. arguta fruit is similar to the commercial kiwifruit, but I find it milder, with less acidic tang. A connoisseur might describe a more sophisticated bouquet of flavors. My first experience with this plant was back in college when a horticulture professor brought a basket of the fruit for the class to sample. I was amazed that the fruit existed-- a bite-sized, thin-skinned version of one of my favorite fruits. I quickly became obsessed with the vine, but I did not encounter it again until I came to the Arnold Arboretum where I got to experience the plant as a whole: the glossy leaves borne on bright red petioles, the exfoliating bark, and the delicate and rather inconspicuous white flowers. Native to northeastern Asia, Actinidia arguta numbers among sixty different species in the genus, but only a handful of these can be grown in colder climates. The common species, A. deliciosa, is only hardy to USDA Zone 8, which means the species cannot be grown in New England, while A. arguta can survive to a remarkable Zone 3. The Arboretum currently holds five additional kiwi species, which all produce edible fruit in various colors and shapes. Despite the taste and ornamental foliage, Actinidia arguta is a fast-growing vine that has escaped from cultivation in western Massachusetts, the New York metro, and northern New Jersey. This complicates any recommendation to introduce the species (which can climb more than thirty feet in a single season) as a more widespread fruit crop, although I have never observed any spontaneous seedlings on the grounds of the Arboretum. Three of our accessions of this species represent wild provenances. A particularly note- worthy plant (accession 905-85*A) is located on the second vine terrace in the Leventritt Garden, twinning up a steel trellis. This accession was received from the Chollipo Arboretum in 1985, which collected the seed on Mount Gaya, in North Gyeongsang Province, South Korea. I'm partial to its sweet wild-tasting fruit and its impressive girth at the base, which sprouts into twisting curls on the trellis. Two additional plants (accession 403-97*B and C) on the upper terrace of the Leventritt represent a wild provenance in Jilin Province, China, where seed was collected from a deciduous mountain forest by the North America-China Plant Exploration Consortium (NACPEC). These two plants have been trained to arch over the path, allowing visitors to view the beautiful structure of the vine from beneath. Buds that produce flowers and fruits occur on the interior portion of the current year's growth--usually obscured beneath the foliage. Actinidia arguta vines are typically dioecious, which means that two vines are needed to produce fruit--one with female flowers, the other with male flowers. The flowers may look very similar, however, because most flowers have both male and female parts, but only the males produce viable pollen and only the females have properly developed structures for receiving pollen and developing fruit. To confuse matters, some plants can produce both male and female flowers, and others have been reported with perfect flowers. The species may even change sexual expression from year to year. The specimens of A. arguta that I have observed at the Arboretum have never been consistent in their fruit production. Hardy kiwi may never become a common fruit crop, and perhaps, given its swift growth and ability to escape from cultivation, it never should. Yet as you stroll through the pathways of the Arboretum, don't forget to stop to investigate our winding vines; you may discover hidden gems nestled beneath the leaves. Rachel A. Brinkman is the assistant manager of horticulture at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23464","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14ebb28.jpg","title":"2018-76-2","volume":76,"issue_number":2,"year":2018,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Exploring the Native Range of Kentucky Coffeetree","article_sequence":1,"start_page":2,"end_page":16,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25646","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060a326.jpg","volume":76,"issue_number":1,"year":2018,"series":null,"season":null,"authors":"Carstens, Jeffrey D.; Schmitz, Andy","article_content":"Exploring the Native Range of Kentucky Coffeetree Andy Schmitz and Jeffrey Carstens M waterway, which has cut a deep ravine through the dry grasslands. Most of the tree species followed this creek, and sure enough, tucked near stands of mesquite (Prosopis glandulosa) and western soapberry (Sapindus saponaria var. drummondii), several coffeetrees were growing. The coffeetrees stood small and stunted--the largest barely exceeding thirty feet--and the pickings were slim, with only a handful of the thick leguminous pods dangling in each tree. We quickly went to work shaking the pods free and recording measurements and habitat data. ALL IMAGES BY THE AUTHORS UNLESS OTHERWISE NOTED orning temperatures clung above freezing when we pulled our minivan onto the dirt roads of Elm Creek Ranch, southeast of Shamrock, Texas. It was March 2015, and we were searching for seed of Kentucky coffeetree (Gymnocladus dioicus). Although we never would have expected to find the species growing in the Texas Panhandle, beyond the range shown on distribution maps, a 2007 herbarium voucher confirmed it was \"locally common\" on the property. The ranch manager, J.C. Brooks, led us to the namesake A native population of coffeetrees (Gymnocladus dioicus) at Elm Creek Ranch, Collingsworth County, Texas, was found beyond the conventionally recognized range for the species. William Carr, J. C. Brooks, and Bob Fulginiti collected the herbarium specimen (TEX 00433298) that first pinpointed this location in 2007. Gymnocladus dioicus 3 herbarium vouchers. At a gas station one afternoon, a man noticed our unusual dash collection and interjected with understandable and friendly curiosity, \"Mind if I ask, what's with the branches in your window?\" Coffeetree Collaboration Our 2015 collecting expedition marked the sixth year of a partnership between the Brenton Arboretum, located in Dallas Center, Iowa, and the National Plant Germplasm System (NPGS) genebank in Ames, Iowa. The partnership has aimed to develop a comprehensive collection of Kentucky coffeetree--sampling populations from across the range of the species, which extends from Ontario through central Arkansas, from west-central Ohio through Oklahoma, along with parts of Kentucky and Tennessee. Although the trees in Texas were scraggly and small, many of the specimens we have seen throughout the years have been impressively grand, measuring well over one hundred feet tall. We think Gymnocladus dioicus should be planted more widely in urban environments. The species has no serious insect or disease Kentucky coffeetree (Gymnocladus dioicus) stands among the largest North problems; it is drought tolerant American members of the bean family (Fabaceae). Andy Schmitz has collected an original copy of this hand-colored engraving by Pierre-Joseph Redout?, along and adaptable to tough soil condiwith other Gymnocladus prints. The engraving first appeared in Henri-Louis tions; moreover, it is exceedingly Duhamel du Monceau's sixth volume of Trait? des Arbres et Arbustes que attractive, with distinctive bark L'on Cultive en France, published in 1815. (even at a young age), interesting Three hours later, the temperature had rocketed compound leaves, and yellow fall color. The to 74?F (23?C), and we drove away almost giddy species should be included among the diverse about collecting Kentucky coffeetree in Texas. tree genera that are used to replace ashes (FraxBy this collection on the fifth day of a nineinus), removed because of emerald ash borer (Agrilus planipennis), and oaks (Quercus), sufday expedition, our van was filling with bags of fering from oak wilt (Bretiziella fagacearum). the beautiful yet odoriferous pods--collected Yet if Gymnocladus is planted more broadly, from sites in Oklahoma and Kansas, in addition we realize that a collection of diverse germto Texas. The dash had become covered with plasm will be needed--both now and far in the stout sticks, which would eventually become MICHAEL DOSMANN 4 Arnoldia 76\/1 ? August 2018 Andy Schmitz stands beneath an exceptional coffeetree in Aurora, New York, which proved to be the largest specimen observed by Carstens and Schmitz over ten years of collections. This cultivated tree measured 110 feet (33.5 meters) tall, 60 feet (18.3 meters) wide, and 57.3 inches (1.4 meters) in trunk diameter at breast height, earning a big tree score of 305 and recognition as a New York state champion. Gymnocladus dioicus 5 future--to make selections adapted to regional climatic conditions and to preserve germplasm for potential reintroduction into the wild. The Brenton Arboretum's first collecting trip occurred in 2008, but plans for the project originated in 2004, as the institution contemplated developing a Nationally Accredited Plant Collection through the American Public Gardens Association's Plant Collections Network (PCN) program. Botanical research had been part of the Brenton's mission since it was established in 1997. After meeting with Mark Widrlechner, horticulturist at NPGS and the PCN's Iowa recruiter, Kentucky coffeetree was determined to be an excellent focus. Gymnocladus dioicus was not currently a PCN collection, and the NPGS had only six viable seed accessions of the species from known wild origins. Moreover, everyone agreed Gymnocladus was definitely underused in urban landscapes. Prior to 2004, the Brenton had eleven accessions of Kentucky coffeetree, but none were wild collected. Andy Schmitz, the director of horticulture, made the Brenton's first wild collection of Gymnocladus in 2008 at Ledges State Park in Boone County, Iowa. He made a few additional collections the following year, but he knew that as the only horticulturist at an arboretum operating under a small budget, additional help was needed to make their future PCN collection a reality. This solidified a long-term relationship between two Iowa horticulturists: Andy Schmitz and Jeffrey Carstens at NPGS. Collecting Seed and Data One of the many benefits of having two institutions striving towards a common goal of preserving genetic plant diversity is the shared work load, especially during the preparation for a collection trip. The success of each trip is largely dependent on the initial identification of specific collecting locations. For the 2015 expedition, for instance, herbarium records and floristic surveys were referenced to identify the anomalous population in Texas, as well as other populations near roadsides and deep canyons, creeks and national battlefields. Further communication with property owners and local botanists--including those who collected the original herbarium specimens--is also beneficial, whenever possible. Local contacts occasionally provide us with GPS coordinates for fruiting specimens, but our efforts typically depend on a pair of highquality binoculars. The characteristic brown pods are easily recognized from considerable distances (even when observed at sixty-five miles an hour) and resemble a flock of blackbirds perched high in the canopy. Thankfully, the fruits are persistent from October through May, allowing us to collect in the winter, when they are highly visible in the leafless canopy. (What other species provides a more than six-month window of fruit senescence?) Collections in early to mid-winter were difficult, however, because the stringy and tough peduncle does not release the fruits as easily, whereas roughly six weeks before bud break, the fruit is easily shaken from the tree. After we have spotted the trees, teamwork makes the seed and data collection easier. Our collection on March 4, 2013, illustrates our basic procedure. On the third day of a nineday expedition through southern Indiana and Kentucky, we were truly looking forward to exploring Griffith Woods, south of Cynthiana, Kentucky, which is known for harboring the world's largest chinkapin oak (Quercus muehlenbergii) within an exceptional oldgrowth savannah. As soon as we pulled into the parking lot, we could see fruiting coffeetrees on the distant horizon. Although we harvested from seven trees, the find of the entire trip was the sixth--discovered some four hours later. This was the second-largest coffeetree we had ever seen, measuring 120 feet (36.6 meters) tall, 46.5 feet (14.2 meters) wide, and 42.0 inches (1.1 meters) in trunk diameter at breast height, and it currently reigns as the Kentucky state champion. To harvest seeds from such a large tree, we used a Big Shot ? line launcher to accurately propel a weighted bag and line over branches high in the canopy. We then used the line to shake pods free. The launcher--a slingshot mounted on an eight-foot pole--has proved its worthiness over and over. On early trips, before using the launcher, we averaged up to two hundred seeds per tree, but now we average six hundred to eight hundred. BOB CUNNINGHAM 6 Arnoldia 76\/1 ? August 2018 Jeffrey Carstens uses a pole-mounted slingshot to collect pods along the North Folk River in Douglas County, Missouri. At right, coffeetree pods at the Brenton Arboretum. The collection time at each mother tree takes around forty-five minutes. Jeff is the \"shaker\" while Andy is the \"gatherer.\" Ideally, enough pods are shaken to the ground to fill one or two five-gallon buckets. While the pods are gathered and packed into labeled sacks, we record GPS coordinates, elevation, associate species, habitat notes, and descriptions of plant health and abundance. Over the years, we expanded our data collection to include height and trunk girth (diameter at breast height) for each mother tree, along with trunk measurements for all woody species over four inches (ten centimeters) within a prescribed area (calculated with a small lens, known as a ten-factor wedge prism). We take herbarium vouchers of branches and fruits, which are later deposited at regional herbaria, and we also photograph each tree in the field. (Back at NPGS, we also scan images of the fruits and seeds for each tree, so that precise dimensions are documented.) Our goal is to capture the potential genetic diversity at every site. Sometimes we collect seed from six to eight mother trees, but this has ranged from two to ten. Sometimes the trees are found less than a few hundred yards apart in a forested area, and other times as much as five miles apart along a river corridor. After we collected from the massive specimen at Griffith Woods, the afternoon sun was setting fast. We never found the record chinkapin oak, because everywhere we turned, massive lookalikes made us freeze in our tracks in admiration. We collected from one other coffeetree--our seventh for the site--and then, as we walked back to the van at twilight, we were startled by a black object moving in the tall grass ahead of us. When our eyes focused, Gymnocladus dioicus 7 we came to realize it was a skunk. Good thing for us (and for the hotel staff), the skunk disappeared without incident. Soon afterwards, several wild turkeys flew overhead, landing in the brush to roost for the night. Observations on Abundance To cover the entire native range of Kentucky coffeetree, we have targeted collection sites approximately seventy-five miles apart within distinct watersheds. We have also aimed to collect from every possible Omernik ecoregion (Omernik, 1987), in an effort to find the best representation of genetic diversity across as many unique habitats as possible. To date, we have made at least one collection from twenty of twenty-two ecoregions within the core range, although small disjunct populations could occur in another twelve. Our first extended collection trip occurred in 2010, when we spent eight days on the road making thirteen collections in six states (Iowa, Missouri, Arkansas, Illinois, Indiana, and Tennessee). On future trips, we usually focused on a single state or a few adjacent states, often targeting two collection sites per day, sometimes as much as one hundred miles apart. On all trips, our work started before sunrise and ended after sunset. Looking at the range map for Gymnocladus dioicus in Elbert Little Jr.'s Atlas of United States Trees, we might assume the species would be well represented near the central part of the range and become scarcer towards the edges, but this assumption does not hold true. Missouri, for instance, is centered within the native range, and approximately 80 percent of the state's Carstens and Schmitz's collection sites are pinpointed on Elbert Little Jr.'s Gymnocladus dioicus range map (USGS, 1999), which has been superimposed over color-coded Level III Omernik Ecoregions (Omernik, 1987). 8 Arnoldia 76\/1 ? August 2018 counties document its presence (Kartesz, 2015). Yet out of nine Missouri populations that we've sampled, finding more than twenty-five trees proved difficult, especially in the southeastern corner of the state--nearly the center of the range. Jeremy Jackson, from the United States Army Corps of Engineers, supplied us with forestry plot data for the Wappapello Lake Project in Wayne County, which demonstrated that only three genetically distinct colonies occurred within the ten-thousand-acre property. Our first-hand observations confirmed this scarcity. This aligns with observations by Gifford Pinchot, the first chief of the United States Forest Service, whose 1907 report suggested that coffeetree was one of the rarest forest trees despite its rather extensive range and that, in large areas within the range, the species was \"entirely lacking or represented only by an occasional individual.\" For this reason, we were especially intrigued by an 1899 report from the geologist Robert Ellsworth Call, which stated coffeetree was \"of very common occurrence\" along aspects of Crowley's Ridge, a geological formation that runs from southeastern Missouri through northeastern Arkansas, paralleling the alluvial plain of the Mississippi River. We wanted to target Crowley's Ridge during our first joint collecting trip, so we spent numerous hours searching the internet and communicating with botanists hoping to pinpoint locations. Only one botanist--a man who had spent more than thirty years studying the area--could tell us of a single population along the ridge. What changed over the past century that has caused the \"very common\" to become rare? Certainly this period coincided with environmental transformations rendered by agriculture, deforestation, grazing, and timber use. Based on our observations, Ken- Coffeetrees proved most abundant on the western edge of its range, where the environment is the most hot and dry. Carstens and Schmitz encountered this specimen growing along a gravel road in Roger Mills County, Oklahoma. Gymnocladus dioicus 9 tucky coffeetree is essentially rare throughout the eastern three-quarters of its native range. While Little's map does not capture this offcentered distribution, general patterns hold true. In 2010, we spent almost a full day scouring areas south and west of Carbondale, Illinois, covering hundreds of miles in and near the LaRue-Pine Hills Research Natural Area, where limestone and sandstone outcroppings tower over the Mississippi River bottoms, yet we only found one lone tree at the base of Fountain Bluff. Our struggle to locate the species in this region aligns perfectly with Little's map, which shows a distribution gap for southern Illinois. In Michigan, on the northern edge of the native range, numerous botanists have kept an eye out for coffeetree due to its rarity. During a seven-day tour of the state in 2016, we were able to locate coffeetree only at locations that had been provided to us and at no additional sites, thus confirming its rarity. In Minnesota, the map shows a few disjunct populations. Our sampling, again dependent on the observations of other botanists, found Gymnocladus dioicus to be infrequent but locally common along watersheds of the Minnesota and Blue Earth Rivers, which once again aligns with Little's map. Compared to other states sampled, Gymnocladus dioicus is quite abundant in Oklahoma and Kansas, where it could be considered a dominant forest species at some sites. Perhaps a combination of historical and current land management practices allow it to sustain and regenerate within these two states. Observations on Habitat When we initially started making seed collections of Gymnocladus near Iowa, we specifically targeted watersheds and bottomlands, as By day nine of the collecting trip through Kansas, Oklahoma, and Texas, Schmitz and Carstens had packed their van with 1,335 pounds of pods and seeds. 10 Arnoldia 76\/1 ? August 2018 In southern Indiana and Kentucky, for instance, Gymnocladus dioicus occurs on upland bluffs and steep slopes with loose soils and occasionally on bedrock. The Loess Hills of western Iowa, which follow the Missouri River, support substantial specimens of G. dioicus on all aspects of their slopes. In Iowa and Illinois, human settlement and introduction of the plow has likely eliminated many G. dioicus in open fields and has left remnants only in areas too wet or steep for modern agriculture. Coffeetree was found in a variety of habitats in Missouri, including moist ravines, dry rocky slopes, and major and minor watersheds. On the western edge of its native range in Kansas and Oklahoma, coffeetree occupies dry ravines and hillsides, open pastures and bedrock. All six sites sampled in Tennessee were collected off cool north- to northeast-facing slopes, showing a definite variance compared to other states. While most sites tend to have rather uniform habitats for all trees sampled, a few possessed specimens growing in remarkably different conditions. Along the Minnesota River in south-central Minnesota, coffeetrees grow on bottomland alluvial soils as well as on threeAndy Schmitz stands beside a multi-stemmed coffeetree on dry bluffs overlooking the Arkansas River in Osage County, Oklahoma. Trees were also billion-year-old granite outcrops, found on the nearby floodplain, demonstrating the remarkable adaptability emphasizing the ability of the speof the species. cies to perform on an extreme specthis is where we found the species naturally trum of harsh growing conditions. Specimens occurring; moreover, this preference for moist at the Tallgrass Prairie Preserve in northeastern locations concurred with descriptions in scienOklahoma demonstrate similar adaptability: tific literature. Yet, over the ensuing years, our we found trees growing on shallow sandstone understanding of the species began to change. outcrops with blackjack oak (Quercus mariWhen we collected in central Michigan in landica) and post oak (Q. stellata), and in moist floodplains with sycamore (Platanus occidenta2016, we noted that even though the majority of lis) and black walnut (Juglans nigra). the specimens were growing in extremely wet Ten years ago, we assumed silver maple (Acer bottomlands, adjacent to major rivers, many saccharinum) would be a common associate, displayed signs of root rot and decline. Whether given our original understanding that Gymnothese Kentucky coffeetrees really preferred to cladus dioicus was a floodplain species, but in be in such wet conditions was questionable. Gymnocladus dioicus 11 Threat of Impermeable Seeds On March 5, 2013, the day after our collection at Griffith Woods, Kentucky, we began before sunrise, heading northeast for the Fleming Wildlife Management Area. This site occupies the eastern edge of the range for Gymnocladus in Kentucky. Scott Freidhof, a wildlife biologist for the area, had provided us with GPS coordinates for coffeetrees there, and this information proved essential. The trees were a three-quartermile hike up to the top of a bluff, and because the forest cover was dense, there was no using binocs to spy pods from afar. With the GPS unit in hand, we attempted to take the most direct route to this localized population and scrambled straight up a steep slope. At the top, pods littered the ground, and we swiftly scooped up twenty-five gallons. The slingshot pole came in handy, not for slinging the throw line but for stringing up the bags of pods like wild game from a hunt. With one end of the pole on each of our shoulders, we made our way down the steep slope, only stumbling a few times on loose rocks or wet oak leaves underfoot. SCOTT FREIDHOF the end, silver maple has been documented at only 12 percent of our sites. Today, our efforts to find coffeetree typically require a watchful eye for common hackberry (Celtis occidentalis), often growing where wet habitat grades into slightly higher elevations or sandier soils. We have observed hackberry--the most common associate--at 62 percent of our collection sites, including on the edge of the Cass River in Tuscola County, Michigan, where we discovered Gymnocladus about seventy miles north of the presumed northern limit for its range. Hackberry and coffeetree regularly share upland locations as well. The common belief that coffeetree is adapted for wet sites in the landscape likely stems from the occurrence of coffeetree amongst bottomland forests of major watersheds in the states of Iowa, Illinois, Minnesota, and Indiana, where sand and loam soils provide adequate drainage. Seeing a strong presence of Gymnocladus in Kansas and Oklahoma, more than any other states, suggests the species is well adapted for dry, hot environments. Because few living animals are known to disperse coffeetree seeds, an abundance of pods can sometimes be found beneath the trees, including here, in Fleming County, Kentucky. Andy Schmitz (left) and Jeffrey Carstens (right) carry a load of pods. 12 Arnoldia 76\/1 ? August 2018 KYLE PORT Why were so many seeds undisturbed on the ground? The Kentucky coffeetree has been referred to as a \"botanical anachronism,\" one that was once dispersed by large prehistoric mammals that are now extinct (Zaya and Howe, 2009). Grinding molars and intestinal juices of the American mastodon (Mammut americanum) may have aided in scarifying coffeetree seeds, and perhaps just as important, these animals would have served as a major dispersal mechanism (Barlow, 2008). Al Fordham, a prominent propagator at the Arnold Arboretum, conducted a germination experiment on Gymnocladus in 1965. He placed three hundred seeds in water, and within the first ten days, thirteen seeds germinated. Fordham suggested \"these, no doubt had fissures in their seed coats.\" Over the next two years, only three more seeds germinated that were submerged in water. Coffeetree seeds are surrounded by a gelatinous material, which may serve as a protective barrier during the early to mid-maturation phases and perhaps later as a reward to animals willing to disperse them throughout the landscape, though which fourlegged critters (if any) now move these fruits remains unclear. Water may be a viable dispersal mechanism. We have observed pods falling into a river, and though able to float for a while, they eventually An open coffeetree pod at the Arnold Arboretum, showing sticky pulp surrounding the seeds (accession 1181-83*A). sink to the bottom, where abrasion provided by gravel and sand in the riverbed may provide the necessary scarification needed for germination. Yet how would such a heavy seed make its way back to shore to even have a chance at sprouting? At many collection sites, two- and threeyear-old seeds can be found on the ground under the canopy of the mother tree, next to rotting pods from prior years. This is what writer Connie Barlow--drawing on the work of ecologists Dan Janzen and Paul Martin--described as the \"riddle of the rotting fruit,\" caused when seed lies in wait for an extinct animal that will never come to carry it away. Our observations recorded little to no regeneration at almost all of our collection sites, illustrating a potential threat for the species within its native habitats. Habitat Loss and Ecological Changes In 2015, the day before we collected Gymnocladus in Texas, we visited a site along the Washita River in Custer County, Oklahoma, where an existing NPGS accession (PI 649669) had been collected in 1993. Steve Bieberich, the owner of Sunshine Nursery in Clinton, Oklahoma, had collected the original accession. We met at his nursery, and he guided us to the site. Even with his help, we were unable to relocate any coffeetrees. By looking at historical photos, we realized a new highway bridge had eliminated Gymnocladus at this location back in 2005. This observation really hit home the importance of our collaborative effort to deposit seeds for long-term preservation. Fortunately, germplasm from this location is currently secure in the NPGS collection, and preserved seed could be used for reintroduction back into the Washita River watershed. In addition to habitat losses like this, habitat modification poses another serious threat. Our main concern lies with Gymnocladus dioicus growing in floodplains of major watersheds. Thirty percent of our collections came from sites like this, but given that 82 percent of these plants were restricted to extremely well-drained soils, a slight change in hydrology (including increased frequency or duration of floods) would significantly impact tree health. At the Michigan sites where we observed serious signs of dieback and root rot, Gymnocladus dioicus 13 Andy Schmitz poses with a grove of coffeetrees in Louisa County, Iowa. Between 2010 (left) and 2016 (right) the trees became overwhelmed with hops (Humulus sp.), causing significant coffeetree mortality that was compounded by increased flooding at the site. we also noted a number of uprooted specimens with debarked trunks lying in water, their root bases sticking eight feet in the air. These collapsed specimens likely resulted from hydrological changes, given that we found fruiting trees on slightly higher ground a few yards away. For a species that comprises only a small percentage of forest canopies, any loss is critical to future conservation. The threat of non-native invasive species (and even aggressive natives) is also significant. Nowhere was this more evident than on the Little Red River near Heber Springs, Arkansas, where back in 2010 it was virtually impossible to not get entangled in Japanese honeysuckle (Lonicera japonica) or bloodied by multiflora rose (Rosa multiflora). We also noted mature coffeetrees acting like trellises for grape (Vitis sp.) and poison ivy (Toxicodendron radicans) resulting in limb decline. That same year, we sampled a site along the Iowa River in Louisa County, Iowa, where we noted more than one hundred genetically distinct trees--the most coffeetrees we had ever seen in a single area. We could have literally collected truckloads of pods. However, when we returned to this location in 2016, we discovered that at least three-quarters of the coffeetrees were dead or in decline after becoming overgrown by hops (Humulus sp.). The area had also flooded numerous times, suggesting a simultaneous and significant change in hydrology. In these real-world situations, even if seeds could germinate or if clonal suckers sprouted, it seems unlikely a new generation of trees would ever have a chance to reach maturity. 14 Arnoldia 76\/1 ? August 2018 Still Collecting now linked to state champion coffeetrees in Kentucky, New York, and Oklahoma. Despite threats to native populations of GymAll told, we've travelled over twenty-five nocladus, we have consistently observed the thousand miles and spent seventy-five days on extreme toughness and adaptability of the spethe road, collecting over a quarter million seeds cies. Not only can it tolerate a wide variety from more than five hundred mother trees. This of soils, but the trees are remarkably durable, has resulted in one of the most comprehensive withstanding severe ice and other unfavorable tree seed collections ever preserved, including weather conditions. When we visited Obion 88 seed accessions in the NPGS (GRIN, Online County, Tennessee, in February 2010, we Database, National Genetic Resources Laboracrossed the mighty Mississippi on the Dorenatory 2018) and 130 accessions of different wild Hickman Ferry and witnessed the aftermath of provenances planted in three informal groupa major storm that had deposited over an inch ings at the Brenton Arboretum. Researchers in of ice a year before our visit. The forest was Ontario, Canada, are already using this expanentangled with downed trees and limbs. Most sive collection for genetic comparisons to their trees were uniformly broken off at a certain threatened native populations. In 2017, a repheight, but fifteen coffeetrees stood undamaged, licated block of 750 trees was planted at the towering above the surrounding mess and callPlant Introduction Station in Ames, representing like beacons to two boys from Iowa. ing fifty wild populations (three mother trees Over the past ten years, our travels have per population and five trees per mother tree), taken us across sixteen states in search of this which in time will help us to learn about polone species: Gymnocladus dioicus. We have linators, growth rates, and hardiness. Yet our spent eight hours on a frigid boat ride on Michifieldwork continues. We still aim to fill gaps gan's Shiawassee River to document the species within ecoregions and sample outlying populafor a new county record. We have tested the tions and unique habitats. The state of Ohio is shocks on our minivan after collecting 1,335 on our hit list because, to date, only one collecpounds of pods during our trip to Texas and tion has been made there. back. We have avoided snow storms in northern Arkansas, but in Kentucky, we collected in the pouring rain. We have worn hip waders too many times to count and used scrapers to clear ice off the inside of the windshield (given the condensation that rises from wet pods stored inside the van). We met an oilman in Oklahoma who just so happened to have a coffeetree pod under the seat of his truck because he wanted someone to identify it for him. We're sure he never thought two guys from Iowa would make the positive determination, let alone two guys from Iowa who just so happened to be on the same backroad Jeffrey Carstens shows the collection of Kentucky coffeetree seeds at the National shaking the exact same pods Plant Germplasm System (NPGS) genebank in Ames, Iowa. Seeds for each individual from a tree. Our names are mother tree are kept in separate packages and are maintained at -18?C (0?F). CRAIG SHIVES Gymnocladus dioicus 15 Andy Schmitz stands at the entrance to the Brenton Arboretum's Gymnocladus collection, which achieved Plant Collections Network accreditation in 2016. Each of the trees (currently 130 wild-collected accessions) has enough space to reach its mature size, while creating an intertwined canopy. More than anything, our adventures have allowed us the opportunity to experience the beauty of this species in the wild and to discover more than what we could ever find in any textbook or publication. In the words of the natural historian William B. Werthner, \"If in your walks through the woods, you chance to come upon a Kentucky coffeetree, count yourself fortunate for it is the rarest of our forest trees.\" And if you do, please give us a call because we would sure like to know its location. References Call, R.E. 1891. The geology of Crowley's Ridge. Annual Report of the Geological Survey of Arkansas for 1889, 2: 1?223. Carstens, J. and A. Schmitz. 2016. Michigan collection trip Gymnocladus dioicus. Technical Report to USDA-ARS Plant Exchange Office, National Germplasm Resources Laboratory, Beltsville, Maryland, 1?42. Carstens, J. and A. Schmitz. 2015. Collection trip report Gymnocladus dioicus. Technical Report to USDA-ARS Plant Exchange Office, National Germplasm Resources Laboratory, Beltsville, Maryland, 1?57. Barlow, C. 2008. The ghosts of evolution: Nonsensical fruit, missing partners, and other ecological anachronisms. New York: Basic Books. Carstens, J. and A. Schmitz. 2014. Collection trip report Gymnocladus dioicus. Technical Report to USDA-ARS Plant Exchange Office, National Germplasm Resources Laboratory, Beltsville, Maryland, 1?18. Baskin, C.C. and J.M. Baskin. 1998. Seeds: Ecology, biogeography, and, evolution of dormancy and germination. San Diego: Academic Press. Carstens, J. and A. Schmitz. 2013. Collection trip report Gymnocladus dioicus. Technical Report to USDA-ARS Plant Exchange Office, National Barlow, C. 2001. Anachronistic fruits and the ghosts who haunt them. Arnoldia, 61(2): 14?21. 16 Arnoldia 76\/1 ? August 2018 Germplasm Resources Laboratory, Beltsville, Maryland, 1?54. Carstens, J. and A. Schmitz. 2010. Collection trip report Gymnocladus dioicus. Technical Report to USDA-ARS Plant Exchange Office, National Germplasm Resources Laboratory, Beltsville, Maryland, 1?18. Fordham, A.J. 1965. Germination of woody legume seeds with impermeable seed coats. Arnoldia, 25(1): 1?8. Herendeen, P.S., Lewis, G.P., and A. Bruneau. 2003. Floral morphology in Caesalpinioid legumes: Testing the monophyly of the \"Umtiza clade\". International Journal of Plant Sciences, 164(S5): 393?407. Harshberger, J.W. 1904. The relation of ice storms to trees. Contributions from the Botanical Laboratory of the University of Pennsylvania, 2: 345?349. Hauer, R.J., Wang, W., and J.O. Dawson. 1993. Ice storm damage to urban trees. Journal of Arboriculture, 19: 187?187. Hoss, G. 2013. Growing difficult hardwoods: Experiences at the George O. White State Forest Nursery. National Proceedings: Forest and Conservation Nursery Associations?2012: 39. Janzen, D.H. and P.S. Martin. 1982. Neotropical anachronisms: The fruits the gomphotheres ate. Science, 215(4528): 19?27. Kartesz, J.T. 2015. The Biota of North America Program (BONAP). North American Plant Atlas. (http:\/\/ bonap.net\/napa). Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2015. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)]. McClain, M.L. 1973. Site preferences and growth responses of Kentucky coffeetree, Gymnocladus dioicus (L.) K. Koch, near the center of its range (doctoral dissertation). Indiana State University, Terre Haute, Indiana. McClain, M.L. and M.T. Jackson. 1980. Vegetational associates and site characteristics of Kentucky coffeetree, Gymnocladus dioicus (L.) K. Koch. Proceedings of the Central Hardwoods Forest Conference, 3: 239?256. Omernik, J.M. 1987. Ecoregions of the conterminous United States. Annals of the Association of American Geographers, 77(1): 118?125. Pinchot, G. 1907. Coffeetree (Gymnocladus dioicus). USDA Forest Service Circular, 91: 1?4. Smith, M.A.L. 1996. Gymnocladus dioicus L. (Kentucky coffeetree). In Y.P.S. Bajaj (Ed.), Trees IV (pp. 194?204). New York: Springer-Verlag Berlin Heidelberg. Smith, W.R. 2008. Trees and shrubs of Minnesota. Minneapolis: University of Minnesota Press. USDA, Agricultural Research Service, National Plant G e r mp l a sm S y ste m. 2 0 1 8 . G e r mp l a sm Resources Information Network (GRINTaxonomy). National Germplasm Resources Laboratory, Beltsville, Maryland. Accessed 1 May 2018 from https:\/\/www.ars-grin.gov\/ USDA, Natural Resources Conservation Service, Soil Survey Staff. 2017. Web Soil Survey. National Soil Survey Center, Lincoln, Nebraska. Accessed 1 May 2018 from https:\/\/websoilsurvey.sc.egov. usda.gov\/ USGS. 1999. Digital representation of \"Atlas of United States Trees\" by Elbert L. Little, Jr. Reston, Virginia: US Geological Survey. Stilinovic, S. and M. Grbic. 1987. Effect of various presowing treatments on the germination of some woody ornamental seeds. International Symposium on Propagation of Ornamental Plants 226: 239?246. Weisehuegel, E.G. 1935. Ger minating Kentucky coffeetree. Journal of Forestry, 33: 533?534. Werthner, W.B. 1935. Some American trees. New York: The Macmillian Company. Zaya, D.N. and H.F. Howe. 2009. The anomalous Kentucky coffeetree: Megafaunal fruit sinking to extinction? Oecologia, 161(2): 221?226. Andy Schmitz has been the director of horticulture\/ general manager of the Brenton Arboretum for twenty years and oversees all horticultural aspects of the 143-acre arboretum, including planting, curating, and maintaining all plant records for over 2,500 woody accessioned plants. He enjoys observing trees in their native habitats while collecting seed to grow trees and shrubs to enhance the arboretum's collections, and his particular focus is on collecting the native Iowa woody plants for their Iowa collection. Jeffrey Carstens is a horticulturist at the North Central Regional Plant Introduction Station in Ames, Iowa, where he currently serves as the curator for the collection of woody and herbaceous plants. He coordinates the NC-7 Woody Ornamental Evaluation Trials and is actively involved with collecting ash (Fraxinus) as part of the National Plant Germplasm System Ash Conservation Project. "},{"has_event_date":0,"type":"arnoldia","title":"Great Wild Gardens: The Story of the Arboretum's Woodlands","article_sequence":2,"start_page":17,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25647","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060a36b.jpg","volume":76,"issue_number":1,"year":2018,"series":null,"season":null,"authors":"Schissler, Danny","article_content":"Great Wild Gardens: The Story of the Arboretum's Woodlands Danny Schissler A Law Olmsted, preserved these masses of native trees, noting their natural beauty and educational potential. \"In no other public garden are there such cliffs or a more beautiful remnant of a coniferous forest,\" Sargent wrote of Hemlock Hill, one of the four main Arboretum woodlands. Of the other areas, he noted that large oaks and other deciduous trees--some more than two-hundred years old, according to his estimate--were valuable for illustrating \"New England trees in their adult state.\" Since Sargent's time, these four areas--North Woods, Central Woods, Hemlock Hill, and DANNY SCHISSLER t the heart of the story of the Arboretum's woodlands lies a tension between the managed and the unmanaged, the natural and the constructed. From the beginning, the Arboretum's woodlands were intentionally excluded from formal development to serve as an aesthetic contrast to the taxonomically grouped collections. The Arboretum's first director, Charles Sprague Sargent, took careful inventory of the remnant woodlands included in the Arboretum's indenture. Rather than clear these areas for collections, Sargent, in concert with the landscape architect Frederick Stands of sweet birch (Betula lenta)--seen here in brilliant fall color--take advantage of canopy gaps on Hemlock Hill. 18 Arnoldia 76\/1 ? August 2018 Four woodlands are traditionally recognized at the Arboretum, shown here from left to right: Peters Hill Woodland, Hemlock Hill, Central Woods, and North Woods. This map was hand-drawn by Jan Tijs Peiter Bijhouwer. Peters Hill Woodland--have come to exemplify the concept of the \"urban woodland,\" providing benefits along with management challenges unique to urban forest fragments. Today, these woodlands provide a naturalistic backdrop to the cultivated collections, offering a sense of spontaneity--whether in a fleeting glimpse of wildlife, discovery of a rare wildflower, or an unexpected encounter with a venerable old tree. Despite the seeming wildness of these areas, the woodlands and the ecosystems they support hardly represent the sort of pristine New England forest we might imagine them to be. On the contrary, they exist at the intersection of the intended and unintended consequences of human decisions--a sustained biological triumph over repeated broad-scale disturbances, creating a colorful mosaic of the native, the non-native, and the outright invasive, while raising questions about the very definition of so-called natural woodlands. Woodland Hill The woodlands inherited by Sargent at the time of the Arboretum's founding in 1872 bore the marks of widespread ecological disturbance, most of it regrown from worn cropland and pasturage. In a 1935 article on land-use history at MODIFIED FROM ARNOLD ARBORETUM ARCHIVES Arboretum Woodlands 19 the Arboretum, research assistant Hugh Raup (who would later be appointed the first plant ecologist on the Arboretum staff) examined deeds of conveyance, records of will, and other historical documents to catalogue the extensive parceling and transfer of properties that would eventually form the Arboretum. From Raup's historical rendering, we know with some certainty that during colonial settlement in the 1700s, the forested portion of the Arboretum's lands befell the same dramatic ecological disturbance as most central New England forests. Rapid deforestation provided fuel and lumber; clear-cut land with fertile soil was cultivated; and upland areas with thin, rocky soils provided pasturage and orchard land. During that period, much of the Arboretum acreage passed through generations of the Davis, Morey, and Weld families. Raup used dendrochronology (a method of tree-ring dating) to show that nearly all of this land--with the exception of the steep slopes of what would later become known as Hemlock Hill--had witnessed the wholesale removal of mature trees. Early on, a saw mill had even been constructed on Bussey Brook. Then, in 1806, a wealthy merchant named Benjamin Bussey began purchasing properties in the area. He consolidated the 20 Arnoldia 76\/1 ? August 2018 rhapsodic reflections inspired by these woodlands, Raup's study of extant trees in 1935 suggests that the oldest hemlocks were scarcely older than thirty when Bussey acquired the hill among his first parcels and, hence, would have been little more than twice that age when Fuller became a frequent visitor. Bussey's stewardship marked a period of rejuvenation for woodlands on the property, yet the ecological succession was nonetheless dictated according to the management practices of this genteel landowner. One local historian, drawing on the memory of older residents in 1897, noted that during Bussey's tenure, woodland paths had been carefully tended all over Hemlock Hill and that an arbor had been erected near the summit, allowing visitors to reflect on the pastoral vista. This aesthetic approach to landscape maintenance was also outlined in Bussey's will, where he dictated that, as long as his family still occupied the land following ARNOLD ARBORETUM ARCHIVES land into an exemplary pastoral estate known as Woodland Hill, on which he would spend his well-earned retirement. Bussey was fascinated by horticultural and agricultural science, and among his three hundred acres of hillsides, meadows, ravines, and brooks, the retired merchant reared merino sheep and cultivated the land through the introduction of novel crop species, trees, and shrubs. Bussey developed his country estate in accordance with the naturalistic English landscaping tradition that had recently permeated American design sensibilities. He targeted areas for reforestation as part of his landscape plan and opened his woodlands, in truly altruistic fashion, to any who wished to escape the bustle and din of nearby Boston. Margaret Fuller and her circle of transcendentalist thinkers visited Bussey's Woods, now known as Hemlock Hill, and she wrote fondly of the soaring hemlocks and pines found along the brook. Despite the Benjamin Bussey's mansion and farm buildings were situated on the eastern slope of Bussey Hill--a short stroll from wooded paths on Hemlock Hill (then known as Bussey's Woods). This inset comes from a large map prepared by Guy Lowell in 1904. At that time, Bussey's home was on property maintained by the Bussey Institution, not the Arboretum. ARNOLD ARBORETUM ARCHIVES Arboretum Woodlands 21 Ernest Henry Wilson captured the tranquil beauty of Hemlock Hill in 1923. The unidentified woman could be Wilson's daughter, Muriel. his death, no trees should be removed, except when necessary \"for the beauty of the groves and the walks.\" Presumably Bussey would have applied a similar approach to other woodlands--or rather thickets of young trees--as he acquired them. The areas now known as Peters Hill Woodlands and the North Woods were primarily ten to fifteen years old when Bussey died in 1842, while the Central Woods--located on rocky soil that was largely unsuitable for agriculture--was between twenty-five and fifty years old. When Sargent took the helm of the Arboretum, his impulse to preserve these woodlands likely spanned the aesthetic and the practical; his views on the multitude of ecological benefits provided by preserved forests--including reduced compaction, mulch creation, windbreaks, and improved soil moisture--are well-captured in his 1875 report, \"A Few Suggestions on Tree Planting,\" prepared for the Massachusetts Board of Agriculture. Sargent initially imagined that the woodlands would serve as plantations for the study of forestry and related sciences. In a letter to Boston's Department of Parks in 1879, Sargent described a \"scientific station\" that would allow for the investigation of \"the best methods of forest reproduction and management\" as well as \"a school of forestry and arboriculture in which special students may ... acquire the knowledge and training necessary to fit them for the care and increase of our forests.\" Eventually, Sargent abandoned this forestry plan, yet he retained the three woodland areas later known as Hemlock Hill, North Woods, and Central Woods, prescribing a basic management regime of occasional thinning--more or less maintaining Bussey's vision for these wooded spaces. Later, a fourth woodland was added with the 1895 annexation of the sixtyseven-acre tract that became Peters Hill. A naturalistic blending of native woodlands and cultivated collections--producing a so-called MODIFIED FROM ARNOLD ARBORETUM ARCHIVES 22 Arnoldia 76\/1 ? August 2018 The undulating borders of Central Woods show the impact of landscape management over five decades. The woodlands were allowed to spread across Conifer Path (originally known as Bridle Path) before undergoing restriction to create space for new accessioned plantings. From left to right: 1937 (or 1938), 1952, 1964, and 1987. landscape effect--formed the foundation for Olmsted's design and left a lasting impact on the institution's identity. Ultimately, the long legacy of human intervention that had shaped the Arboretum's woodlands would continue into the twentieth century and well beyond Sargent's time, as staff members grappled with a succession of natural and unnatural disturbances in these areas. Managed Succession Among the Arboretum woodlands, Hemlock Hill most clearly shows the ongoing process of human intervention. On a cold September evening in 1938, a four-day rainstorm crescendoed across New England. Violent wind gusts buffeted forests south of Boston, and the Blue Hills Observatory recorded hurricane speeds of over 150 miles per hour. At the Arboretum, staff members hunkered down in the darkness of the Administration Building, listening to the creaks and groans of the trees. The worst of the storm lasted only a few hours. The next morning, staff awoke to a grim scene. The Arboretum suffered greatly: over fifteen hundred trees had been claimed by the winds. Much of the damage befell the Arboretum's woodland areas, including Hemlock Hill, where at least four hundred native hemlocks (Tsuga canadensis) lay in splinters. Arboretum staff responded to this cataclysm by planting hundreds of hemlocks in their place, some as large as six feet tall. The storm would prove to be the most destructive in the recorded history of New England, just one in a series of events that transformed the Arboretum's natural woodlands--its marks still visible today. Yet in many ways, the natural history of Hemlock Hill, and the Arboretum woodlands in general, has been a story of ongoing landscape management. Without human intervention, ecosystems respond to disturbances like hurricanes, fire, and even secondary regrowth after agricultural land is abandoned, through the process of succession-- or the gradual change in species structure in an ecological community. Since the beginning of this successional process for the Arboretum woodlands, starting when Bussey set aside reforestation land and allowed seedling thickets to become established, this gradual change has been continually manipulated, especially in response to large-scale disturbance like the hurricane. This management, of course, raises questions about the very conception of natural succession and whether strategies often intended to contribute to (and perhaps simply expedite) these ecological changes are, in fact, additional forms of disturbance. In 1930, nearly a decade before the hurricane Today, the control of invasive plant species is caused ecological upheaval across New Engoutlined as an ongoing objective in the Arboreland, Arboretum botanist Ernest Jesse Palmer tum's Landscape Management Plan, although presented an extensive survey of the Arboremany of these interventions are conducted on tum's spontaneous flora, cataloging biodiversity an ad hoc basis, given that most horticultural throughout much of the grounds--including its resources are invested in the more manicured woodlands. Alongside his thorough inventory portions of the living collections. Nonetheof each area of the living collections and the less, occasional efforts have been devoted to underlying geology of the landscape, Palmer this end. Peters Hill Woodland, for instance, hinted ominously at the effects of aggressive was subject to a three-year project conducted exotic plants on native flora. His account is by the Hunnewell interns, starting in 2008, particularly notable for its description of the with the last two years focused primarily on colonization of highly disturbed areas, such as removing woody plants like cork tree and casthe abandoned quarry south of Bussey Street, tor aralia (Kalopanax septemlobus), which by an \"uncommon\" assemblage of herbaceous had escaped from the surrounding collections. weedy species like green foxtail (Setaria viridis), Control of the botanical composition of urban black nightshade (Solanum nigrum), and comwoodlands--particularly those in close proxmon vetch (Vicia sativa). These species, notaimity to historically disturbed areas--is often bly absent from his inventories of the diverse costly, however, requiring horticultural care and richly populated woodland areas, had only be diverted from the accessioned collections. begun to take hold on the grounds. The time Moreover, the management of invasive spebetween Palmer's and ours marks an ecological cies using mechanical and chemical methods transition for many of the Arboretum's natural raises questions about the very idea of preservareas, with the slow creep of invasive plants ing ecosystem processes, further muddying our gradually shifting the compositions of species understanding of how landscapes continually among these woodland fragments. disturbed by human intervention could be conMost of the first weedy species to show up strued as natural. in New England arrived with European settlers Introduced insects and pathogens have also beginning in the seventeenth century. Wellinspired radical management changes in the adapted to continually disturbed conditions, many of these species established themselves in parts of the Arboretum. A second wave of non-native introductions arrived on a network of exploration and plant trade connecting Western nurseries and botanical institutions with East Asia beginning in the 1860s, resulting in the rapid importation of thousands of potentially invasive species. Through its legacy of collection and distribution of exotic plants, the Arboretum played its part in popularizing many of these species, such as Oriental bittersweet (Celastrus orbiculatus) Oriental bittersweet (Celastrus orbiculatus) twines atop castor aralia (Kalopanax and Amur cork tree (Phelloden- septemlobus). Both are abundant escapees from the Arboretum's cultivated collection, observed here in Peters Hill Woodland. dron amurense). JONATHAN DAMERY Arboretum Woodlands 23 24 Arnoldia 76\/1 ? August 2018 up to fill canopy gaps. In 2006, the Arboretum also planted sapling oaks (Quercus montana, Q. coccinea, Q. velutina), shagbark hickories (Carya ovata), and sugar maples (Acer saccharum) on the southeast side of Hemlock Hill. To echo Palmer's observations from 1930, \"The line between Nature's great wild gardens and those planted and tended by man is not a hard and fast one ... Nature herself is the builder if not the designer, guided only by man's selection and aid in planting, pruning and cultivating the things he deems most desirable.\" Across its rocky terrain, Hemlock Hill bears the marks of past attempts to preserve what Sargent had once deemed \"the great natural feature of the Arboretum\" through generations of stewardship. While a walk in its cool and shady understory may mentally transport us to the \"primeval\" New England forest that even RICHARD SCHULHOF woodlands--a point illustrated by the arrival of hemlock woolly adelgid (Adelges tsugae) at the Arboretum in 1998. After the initial discovery of this destructive pest on Hemlock Hill, a substantial effort on the part of the Arboretum's horticulture and curation staff culminated in the accessioning of over nineteen hundred existing trees (some nearly two hundred years old), allowing for the close monitoring of the spread of adelgid and its impact on the hemlock population. Today, the remaining mature hemlocks--many of them originally planted in response to the destruction of the Hurricane of 1938--owe their survival to annual treatment with a soil- and trunk-injected insecticide, imidacloprid. Where mature trees have fallen or been removed, dozens of recently planted Chinese hemlock (Tsuga chinensis)--naturally resistant to the ravages of the adelgid--reach Arboretum arborist Robert Ervin holds a branch from an adelgid-infested hemlock (notice the cotton-colored egg masses), which was felled within a research plot on Hemlock Hill in early 2005. PHOTOS BY DANNY SCHISSLER Arboretum Woodlands 25 Canada mayflower (Maianthemum canadense) is an abundant spring ephemeral on Hemlock Hill, while spotted cranesbill (Geranium maculatum) is more commonly observed in Peters Hill Woodland. Palmer envisioned there, the turbulent history of this forest fragment and its resulting character is perhaps the most challenging to our notion of what constitutes a natural woodland. Ecology of the Urban Woodland Natural or not, the Arboretum's woodlands support a great deal of biodiversity. In contrast to the cultivated collections, the successional composition, varied topography, and increased leaf litter and woody debris of these areas provide suitable habitats for a variety of native and non-native species. The woodlands harbor a variety of deciduous hardwoods, conifers, shrubs, herbaceous species, ferns, mosses, and fungi. The woodlands also provide habitats for a range of fauna that often avoid open forests and humans. Snags--dead trees that remain standing--and decaying holes in trunks provide shelter for cavity-nesting birds. Tall trees with dense canopies offer nesting opportunities for larger birds of prey. In the shady understory, reptiles and amphibians make homes among the leaf litter and decaying logs. Wild turkeys forage for acorns and nuts from beeches and hickories. A variety of mammals--coyote, deer, foxes, rabbits, raccoons, opossums, squirrels, chipmunks, voles, and field mice--utilize the Arboretum woodlands. In addition to supporting biodiversity, these woodlands provide a range of ecosystem services that benefit outlying collections. Given the Arboretum's location in a densely populated urban environment, the entire landscape faces an exceptional set of disturbance and climatological factors. The constant pressure of competing species, exotic wildlife, and invasive pests and pathogens is compounded by elevated air and soil temperatures, carbon dioxide, ozone and nitrogen levels, decreased humidity and water availability, soil compaction, and the presence of pollutants. The preservation of urban forests combats these factors by promoting soil building and moisture retention, erosion prevention, temperature control, and carbon sequestration. As Sargent had once envisioned, the Arboretum's woodlands complement the surrounding cultivated collections aesthetically, as part of a naturalistic landscape design, and ecologically, as a buffer against the often harsh conditions of the urban environment. While the Arboretum's woodlands never became the forest plantations for the study of continues on page 30 26 Arnoldia 76\/1 ? August 2018 PHOTOS BY DANNY SCHISSLER Woodland Sketches An abundance of fungi occur in the understory of the North Woods, where fallen trees provide habitat. From left to right: shaggy mane (Coprinus comatus) and yellow orange fly agaric (Amanita muscaria var. formosa). North Woods North Woods (2 acres) is situated along eskers that overlook the Leventritt Shrub and Vine Garden. The Arboretum acquired the westernmost part from the Adams Nervine Asylum in 1926, but the remainder has been part of the Arboretum since its founding. The eastern part of North Woods has diminished over time. This area is also home to quite a few non-native species that likely escaped from the cultivated collection, including cork tree (Phellodendron amurense), Korean mountain ash (Sorbus alnifolia), and Oriental bittersweet (Celastrus orbiculatus). Geology and soils: Higher pH soils (A horizon: 4.21; B horizon: 4.47) than Hemlock Hill and Central Woods; glacially deposited eskers underlain by gravel and other sediment; groundcover mostly of deciduous leaf litter. Mid and overstory: Dominated by sugar maple (Acer saccharum), with an abundance of sweet birch (Betula lenta); interspersed with white oak (Quercus alba) and shadbush (Amelanchier arborea). Understory: Woody taxa include many escaped species from nearby collections such as sapphire berry (Symplocos paniculata), euonymus (Euonymus spp.), honeysuckle (Lonicera spp.), linden (Tilia spp.), and zelkova (Zelkova spp.); herbaceous groundcover includes sedge (Carex spp.) and aster (Symphyotrichum spp.). Wildlife species of note: Great crested flycatcher (Myiarchus crinitus), eastern wood-pewee (Contopus virens), wood thrush (Hylocichla mustelina), red-eyed vireo (Vireo olivaceus), ovenbird (Seiurus aurocapillus), black-throated blue warbler (Setophaga caerulescens), black-throated green PHOTOS BY DANNY SCHISSLER Arboretum Woodlands 27 White trillium (Trillium grandiflorum) and common milkweed (Asclepias syriaca) are among the many wildflowers found in the Arboretum's woodlands. warbler (Setophaga virens), black-and-white warbler (Mniotilta varia), eastern screech owl (Megascops asio), and eastern red-backed salamander (Plethodon cinereus). Central Woods Central Woods (6.5 acres) was maintained as pastureland before a period of regrowth beginning in the 1790s. This woodland is favored by wildlife species that prefer dense, mixed forests. While this woodland has been relatively undisturbed, containing few non-native species in comparison to other areas, the dominance of eastern white pine (Pinus strobus), a pioneer species, is the result of formal clearing in some areas several decades ago. Geology and soils: Low pH soils (A horizon: 3.6; B horizon: 4.09) underlain by outcroppings of Roxbury conglomerate in many areas; heavy cover of duff and leaf litter compared to the other Arboretum woodlands. Mid and overstory: Primarily dominated by eastern white pine, red oak (Quercus rubra), and white oak, with stands of immature eastern white pine and American beech (Fagus grandifolia). Understory: Dominated by lowbush blueberry (Vaccinium angustifolium) and huckleberry (Gaylussacia spp.). Wildlife species of note: Red-breasted nuthatch (Sitta canadensis), yellowbellied sapsucker (Sphyrapicus varius), pine siskin (Spinus pinus), common redpoll (Acanthis flammea), purple finch (Haemorhous purpureus), red- and white-winged crossbill (Loxia curvirostra, L. leucoptera), great horned owl (Bubo virginianus), and coyote (Canis latrans). DAVID CAPPAERT, BUGWOOD.ORG 28 Arnoldia 76\/1 ? August 2018 The Arboretum's woodlands provide habitat for a range of fauna, and some, like the redbacked salamander (Plethodon cinereus), are found almost exclusively within these areas. Hemlock Hill Hemlock Hill, the largest Arboretum woodland, occupies 22 acres. It has had a complex history of disturbance, including the 1938 hurricane and arrival of the hemlock woolly adelgid. This woodland is home to a number of unique birds, amphibians, ferns, and herbaceous perennials that prefer the shady understory of dense forestland. Prominent non-native plants include glossy buckthorn (Frangula alnus), castor aralia (Kalopanax septemlobus), mountain ash (Sorbus spp.), and hawthorn (Crataegus spp.). Geology and soils: Low pH soils (A horizon: 3.75; B horizon: 4.19); steep rock outcroppings on northeast side; pit-and-mound formations formed by downed trees throughout. Mid and overstory: Heavily dominated by eastern hemlock and some red oak, along with stands of eastern white pine and sweet birch succeeding mature trees. Understory: Dominated by Canada mayflower (Maianthemum canadense), wild sarsaparilla (Aralia nudicaulis) and hay-scented fern (Dennstaedtia punctilobula), with shadbush (Amelanchier arborea) and mapleleaf viburnum (Viburnum acerifolium). Wildlife species of note: Red-breasted nuthatch, pine warbler (Setophaga pinus), black-capped chickadee (Poecile atricapillus), tufted titmouse (Baeolophus bicolor), red-tailed hawk (Buteo jamaicensis), Cooper's hawk (Accipiter cooperii), Virginia opossum (Didelphis virginiana), eastern red-backed salamander, and northern dusky salamander (Desmognathus fuscus). Peters Hill Woodland The 2.5-acre woodland on the eastern slope of Peters Hill was the subject of the Hunnewell intern project for three years (2008?2010). In 2008, the intern class surveyed the vegetation and came up with management recommendations pertaining to invasive species removal, which the following two classes carried out. Peters Hill is the most species-rich of the woodlands and provides space for the greatest number of non-native species, most notably cork tree, crabapple (Malus spp.), hawthorn, and Korean mountain ash. PHOTOS BY DANNY SCHISSLER Arboretum Woodlands 29 False Solomon's seal (Maianthemum racemosum) and wood anemone (Anemone quinquefolia) are among the spring ephemerals observed in the Arboretum's woodlands. Geology and soils: Comparatively high pH soils (A horizon: 4.28; B horizon: 4.51) due to a lack of conifers; steep slopes forming a wet ravine that provides water throughout most of the year; heavy presence of woody debris, duff, and leaf litter. Mid and overstory: Dominated by red oak, followed by sassafras (Sassafras albidum), black oak, cork tree, yellow birch (Betula alleghaniensis), castor aralia, and a variety of other native and non-native hardwoods. Understory: Dense understory with at least twenty-three woody species in the sample plot; abundance of native and non-native saplings, primarily cork tree; many common herbaceous perennials. Wildlife species of note: Great crested flycatcher, scarlet tanager (Piranga olivacea), rose-breasted grosbeak (Pheucticus ludovicianus), black-throated green warbler, eastern wood-pewee, wood thrush, chimney swift (Chaetura pelagica), common nighthawk (Chordeiles minor), and common garter snake (Thamnophis sirtalis). Survey methods Woody flora was documented in a 2017 survey, based on two randomly assigned ten-meter-radius circular plots within each of the four woodlands. In Peters Hill Woodland, only one circular plot was examined along with a recreated transect first studied by the Hunnewell interns, class of 2008. In addition, each study area was sampled as part of the 2017 landscapewide soil survey. Ten auger samples, separated into A and B horizons, were taken within each of the four study areas and composited, producing one A- and one B-horizon sample for each natural land. These samples were sieved, air-dried, and sent to the University of Massachusetts for analysis. 30 Arnoldia 76\/1 ? August 2018 continued from page 25 lands. Data acquired through these projects can help researchers understand the response of urban woodlands and their inhabitants to a changing climate. Resistance of Nature Since the time of Benjamin Bussey and the wayward philosophers for whom he opened his lands, the forest fragments now situated on the Arboretum's grounds have offered a space for rejuvenation and a retreat from the hum of city life. Sargent and Olmsted--both profoundly influenced by an English tradition of naturalistic park design--incorporated these woodlands as a visual backdrop for the accessioned plant collections, adapting land that Olmsted described in a 1880 letter to author Charles Eliot Norton as largely unfit for cultivation. Today, the Arboretum's woodlands provide visitors a sense of tranquility and privacy JAY CONNOR arboriculture that Sargent had imagined, the unique ecosystems of these areas have served researchers working across a range of scientific disciplines. Sheltered habitats situated among woodland microclimates--such as the shady, moist understory of Hemlock Hill--have offered opportunities to study native salamander species. Mature trees in Central Woods are used in climatological and phenological studies. Peters Hill Woodland, along with the \"urban wild\" of Bussey Brook Meadow, present unique successional models of minimally managed urban vegetation and the ecosystem services provided by cosmopolitan assemblages of species found in such areas. Most recently, for mer Living Collections Fellow Jenna Zukswert collaborated with other staff members to conduct an Arboretum-wide survey of soils and species composition within the wood- The Arboretum woodlands--all visible from this springtime vantage in 2005--record a long trajectory of landscape management practices within a highly modified urban environment. Arboretum Woodlands 31 often missed among the more open and ordered character of the cultivated collections. Here, the allure of wildness and the excitement of spontaneity play out in chance encounters with the seemingly natural. But the character of such spaces begs the question: what is truly natural in an era of accelerated ecological upheaval? What role do such spaces--shaped continuously by the interplay of environmental stochasticity and human impulse--play in the Arboretum landscape today? And finally, how might we manage these spaces to reap the spiritual and ecological benefits they provide, while acknowledging the realities of our rapidly changing urban environments? In the words of Palmer, \"There is a constant effort of Nature to reassert her sway and reclaim for herself the areas that men have planted. Even in the best kept gardens this jealous resistance of Nature is not entirely overcome.\" At the Arnold, the genius of Sargent and Olmsted's collaborative vision lives on in these naturalistic, if not entirely natural, interstitial spaces between the cultivated and the wild--not only in their physicality, but in the way they touch our primal selves, helping us forget, if only for a moment, that we're walking in a garden. Olmsted, F.L. 1982. To Charles Eliot Norton, 5 May 1880. In R.C. Wade (Ed.), The papers of Frederick Law Olmsted: parks, politics, and patronage (pp. 463?465). Baltimore: The Johns Hopkins University Press. References Whitcomb, H.M. 1897. Annals and reminiscences of Jamaica Plain. Cambridge: Riverside Press. Arnold Arboretum of Harvard University. 2011. Landscape management plan (3rd edition). Retrieved from https:\/\/www.arboretum.harvard.edu\/ wp-content\/uploads\/AA_LMP_Summary.pdf Arnold Arboretum of Harvard University. 2007. Cultural resource management plan. Arnold Arboretum Horticultural Library, Harvard University. Bussey, B. 1859. Will of Benjamin Bussey of Roxbury. Boston: J.H. Eastburn's Press. Del Tredici, P. 2010. Spontaneous urban vegetation: reflections of change in a globalized world. Nature + Culture 5(3): 299. Hay, I. 1995. Science in the pleasure ground: a history of the Arnold Arboretum. Boston: Northeastern University Press. Hay, I. 1994. George Barrell Emerson and the establishment of the Arnold Arboretum. Arnoldia 54(3): 12?21. Mathewson, B. 2007. Salamanders in a changing environment on Hemlock Hill. Arnoldia 65(1): 19?25. Palmer, E.J. 1930. The spontaneous flora of the Arnold Arboretum. Journal of the Arnold Arboretum 11(2): 63?119. Raup, H.M. 1935. Notes on the early uses of land now in the Arnold Arboretum. Bulletin of Popular Information (Arnold Arboretum, Harvard University) 3(9?12): 41?74. Sargent, C.S. 1875. A few suggestions on tree-planting. Twenty-Third Annual Report of the Secretary of the Board of Agriculture. Boston: Wright and Potter, State Printers. Sargent, C.S. 1922. First fifty years of the Arnold Arboretum. Journal of the Arnold Arboretum 3(3): 127?171. Sargent, C.S. 1880. To the Board of Park Commissioners, City of Boston. Fifth annual report of the Board of Commissioners of the Department of Parks for the City of Boston for the year 1879 (pp. 21?22). Boston: Rockwell and Churchill, City Printers. Sargent, C.S. 1885. To Frederick Law Olmsted, 5 February 1885. Charles Sprague Sargent (1841?1927) papers, Arnold Arboretum Horticultural Library, Harvard University. Schulhof, R. 2008. Ecosystems in flux: the lessons of Hemlock Hill. Arnoldia 66(1): 22?28. Wilson, M.J. 2006. Benjamin Bussey, Woodland Hill, and the creation of the Arnold Arboretum. Arnoldia 64(1): 2?9. Many thanks to the following staff members at the Arnold Arboretum for offering their knowledge of the Arboretum's woodlands: Rachel Brinkman, Ana Maria Caballero McGuire, Jonathan Damery, Peter Del Tredici, Michael Dosmann, Andrew Gapinski, Wes Kalloch, Jim Papargiris, Lisa Pearson, Sue Pfeiffer, Kyle Port, Kathryn Richardson, Nancy Sableski, Stephen Schneider, and Mark Walkama. Special thanks to Jenna Zukswert for her survey work, data analysis, and contributions to this article. Daniel Schissler is Project Coordinator at the Arnold Arboretum, a for mer research assistant in the Friedman Lab, and a former Isabella Welles Hunnewell horticultural intern, class of 2016. He holds an MFA in photography from Massachusetts College of Art and Design and a BA in architectural studies and archeology from Tufts University. "},{"has_event_date":0,"type":"arnoldia","title":"Hickory Fever: Doing Taxonomy by Mail","article_sequence":3,"start_page":32,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25648","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d060a76f.jpg","volume":76,"issue_number":1,"year":2018,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"Hickory Fever: Doing Taxonomy by Mail Jonathan Damery B otany in the early years of the Arnold Arboretum required a good postman. Boxes of photographs and herbarium specimens passed back and forth on the railroad. Taxonomic questions would follow in letters, along with requests for more specimens (and usually more again). Charles Sprague Sargent, the founding director of the Arboretum, famously obsessed over the taxonomy of hawthorns (Crataegus). His work was comprehensive and exhaustive, leaving no leaf or flower unturned, and as such, his letters are filled with requests for specimens, fruits, and field descriptions of these small, confusing trees. Yet if hawthorns were first on Sargent's mind, hickories (Carya), the prominent forest trees of eastern North America, were not far behind--often mentioned in the same burst of typewriter keystrokes. Of course, Sargent travelled widely and frequently to study plants in the field--camping on mountainsides, riding in motorcars on dirt roads--but for a project like the Silva of North America, a fourteen-volume work on tree species native to the United States that was published between 1891 and 1902, Sargent needed assistants far and wide. The same was true for subsequent projects that aimed to disentangle specific taxonomic problems, like his synoptic treatment of North American hickories published in the Botanical Gazette in 1918. While Sargent worked on these projects--studying specimens and hand-written field descriptions at his desk on the third floor of the Arboretum's administration building--it must have felt like reconstructing a crime scene from several states away, years after the fact, with only a team of freelance detectives who could occasionally be marshalled (when time and finances permitted) to search for evidence and knock on doors to interview witnesses. In a long and detailed letter to Thomas Grant Harbison, one of his most reliable field collectors in the southeast, Sargent professed immense confusion when it came to the hickories. \"It begins to look as if all the characters on which we have been trying to base species are giving out,\" he lamented in 1914, after providing several pages of notes on specimens Harbison had collected. \"I think that ... the same species may bear globose and oblong nuts, that the leaves may or may not be pubescent, and that the bark may vary enormously according to situation.\"1 ALL IMAGES FROM ARNOLD ARBORETUM ARCHIVES UNLESS NOTED Hickory Taxonomy 33 Above and facing page: Charles Sargent studied hickory (Carya) taxonomy with support of a large network of field correspondents, including the journalist Charlie C. Compton (\"Miss Compton\") in Natchez, Mississippi. \"No. 22 is still puzzling,\" Sargent wrote in 1913, responding to this set of Compton's photographs and field notes. \"Thanks to your energy and intelligent zeal we shall get to the bottom of this business sooner or later.\" Sargent later determined this tree, No. 22, was a black hickory (Carya texana). Evidence for solving these taxonomic mysteries could be frustratingly scant. Even if someone collected herbarium specimens in the middle of the growing season, Sargent would send them back to collect again in the fall. Specimens in hand often only confirmed that others were needed. When he received fruits from a hickory that Ernest Jesse Palmer had collected in Noel, Missouri, in 1915, Sargent told him it was \"one of the most remarkable of all your Hickories,\" yet the fruit had only wetted his desire to know more. \"Will you tell me something about this tree, its size, place of growth, character of the bark, or anything else you may know about it? I have never seen any fruit like this.\"2 Four years later, Palmer would send grafted material from the same tree--now considered a mockernut hickory (Carya tomentosa, accession 8014*A)--which still grows in the Arboretum's hickory collection. Although spindly (much smaller than expected of a centenarian overstory species), the tree offers a robust reminder of the correspondence needed to conduct this kind of taxonomic inquiry. Special Agents Sargent's crew of field correspondents began solidifying long before his interest in hickories. His first concerted research project was the Report on the Forests of North America (Exclusive of Mexico), an ambitious opening salvo launched as part of the 1880 United States Census, which aimed to describe and map the composition of forests across the country. Sargent 34 Arnoldia 76\/1 ? August 2018 Sargent's Report on the Forests of North America (Exclusive of Mexico), published in 1884, included the first distribution maps for major North American tree genera, which were prepared by Andrew Robeson. Rather than presenting the distribution of individual Carya species on this map, the green shading suggests species richness--or the number of Carya species believed present at any particular site. The greatest density (eight species) was recorded in western Arkansas, the future location for significant collections by Sargent and his correspondents. Hickory Taxonomy 35 logged significant miles to research the project himself--notably botanizing forests in Utah, California, and Oregon--but to complete such a wide-ranging project, he needed colleagues that ranged just as far. Several botanists were officially enlisted as \"special agents\" for the four-year project, but others became informal collaborators.3 A number of the oldest hickories still growing at the Arboretum arrived due to the census project, including an exceptional specimen of pecan (Carya illinoinensis, accession 12913*A), tucked in the back corner of the hickory collection, where its straight trunk lofts the canopy nearly one hundred feet high. Fruit for this accession arrived from the ornithologist Robert Ridgway, the first full-time curator of birds at the United States National Museum, who had collected the material near Mount Carmel, Illinois, in 1882. Although Ridgway was based in Washington, DC, he continued to study the landscape of Illinois--his home state--while he prepared a two-volume treatise on the birds of the state. Ridgway was an unofficial census correspondent. Yet his research on woodlands in southern Illinois (and adjacent portions of Indiana) was so rigorous--far surpassing the needs of the census--that Sargent encouraged him to publish his findings independently. The article ran forty pages in the Proceedings of the United States National Museum, published in 1882, the same year the Arboretum received seed shipments from Ridgway. In the report, Ridgway described the pecan as \"one of the very largest trees of the forest\" with a canopy that often \"reared conspicuously above the surrounding tree-tops, even in a very lofty forest,\" and he noted that one tree (unfortunately measured after it had been felled) had been documented with a canopy 175 feet (53 meters) high and a trunk diameter of 5 feet (1.5 meters).4 The pecan in the Arboretum collection, while not yet that size, suggests this pedigree. A stand of nine shellbark hickories (Carya laciniosa, accessions 12898 and 20094) that grow in park-like planting atop Valley Road also arrived in 1882 from another census correspon- An unattributed Garden and Forest editorial from 1889 suggested that hickories \"are the despair of people who expect to be able to fit exactly every plant they encounter with the printed description of it in some book.\" This supplementary illustration of a shellbark hickory (Carya laciniosa) was based on a Robert Ridgway photograph from southern Indiana. dent. George Washington Letterman had been enlisted as an official special agent to study the forests west of the lower Mississippi River, although the hickories were collected near his home in Allenton, Missouri, about thirty miles west of Saint Louis. Letterman was a school teacher and scheduled fieldwork around his classroom duties. In a humorous note to George Engelmann, a prominent botanist who was a physician and a close mentor to Sargent, Letterman alluded to this time constraint in April 36 Arnoldia 76\/1 ? August 2018 1881. Because so many students had the measles that spring, he suggested he might cancel classes and gain an unexpected week for census fieldwork. In the same note, Letterman also described the perplexities of hickories. \"It seems that the hickory nuts puzzle every body, especially those who have not been able to see the trees in all stages of growth year after year,\" he wrote, referring to an inquiry from the Illinois botanist George Vasey. \"Don't you think that something better than what the books now contain on the subject should be given to botanists? In case you undertake to revise the genus, I should be glad to procure all the material obtainable here for you.\"5 Engelmann responded with a brief postcard: \"Too early to work up Carya, but we must go on gathering material.\"6 The epistolary trail with Engelmann ends there, although Sargent, writing two decades later, recalled that Letterman made substantial collections for Engelmann near Allenton and that those collections included \"many notes on the Oaks and Hickories.\"7 Entirely Overlooked For his part, Sargent didn't seem espe- Above and facing page: \"This particular form of Hickory is quite new to cially interested in the hickories dur- me and I am anxious to have further information about the tree,\" Sargent ing the census years, and he wouldn't wrote to Bryant K. McCarty--a pineapple farmer in Saint Lucie County, begin to wade into the subject until an Florida. The herbarium specimen had been forwarded in 1911 from Robert Morris, Sargent's collaborator on hickory matters. Sargent later named the 1889 Garden and Forest article, where species Carya floridana--the scrub hickory--and Charles Faxon provided he attempted to parse out whether the the first illustration. genus should be called Carya, Hicoria, or Hicorius, opting for the final option.8 the different species, which is hardly surprising, An unattributed editorial ran after this nomensince botanists themselves are often perplexed clatural treatment, celebrating the hickory as over questions concerning the proper limita\"purely an American tree,\" given that none of tions of these species.\"9 the Asian species were known to Western botaEven so, when Sargent covered the genus in nists at the time. As the \"conductor\" for the the seventh volume of Silva of North America, magazine (essentially the publisher), Sargent published in 1895, he sounded little taxonomic must have conceded the general points, includalarm, although he footnoted a new variety of ing, quite notably, an admission of taxonomic pignut hickory (what he called Hicoria glabra confusion. \"More Americans know the Hickvar. villosa), based on a tree Letterman had docory-tree when they see it than any other of our umented in Allenton, and he offered passing trees,\" the author wrote. \"That is, they know descriptions of several unnamed hybrids. Yet, generally, the Hickory, without distinguishing hickories weren't alone in escaping Sargent's ARNOLD ARBORETUM HERBARIUM Hickory Taxonomy 37 ARNOLD ARBORETUM HERBARIUM 38 Arnoldia 76\/1 ? August 2018 Hickory Taxonomy 39 According to Sargent's correspondence records (which become more consistent in the Arboretum archive in 1902 when he began saving carbon copies of his typewritten letters) his interest in hickories began gaining traction in 1908. That fall, around the time that hickory fruits would be ripening, he received a letter from a physician-turned-botanist named Robert T. Morris, who inquired about Carya buckleyi (now considered C. texana, the black hickory). Although the two men had corresponded about hickories the year before, Morris's question about the black hickory seemed to awaken Sargent's curiosity. \"I confess that I, as well as all other botanists in recent years, have entirely overlooked this tree,\" Sargent wrote back, referencing the taxon at large, rather than an individual plant. \"The name does not appear in my Silva for some unaccountable reason as I was familiar with the paper [in the Proceedings of the Academy of Natural Sciences of Philadelphia] where it was first described ... I shall be very glad of some of the nuts if you can spare them for me.\"10 Morris had obtained a letter about the species from a grape breeder in northeastern Texas named Thomas Volney Above and facing page: After Sargent visited Van Buren, Arkansas, in Munson. Sargent wrote to Munson March 1909, his local collaborator, George Brown, collected flowering herimmediately, even before responding barium specimens that Sargent would use to describe a new hybrid: Carya to Morris, and offered to trade a selec? brownii. Charles Faxon provided the first illustration of this hybrid. tion of Chinese grape seedlings (grown taxonomic scrutiny; his research interests had from Ernest Henry Wilson collections) for fruit just begun shifting from nomenclatural synand herbarium specimens from the hickory.11 thesis to novel taxonomy. Over the preceding This exchange proved successful. Within three years, Sargent had described as few as twentyweeks, Munson had already shipped the specione new taxa for an assortment that included mens, along with a list of grape species he was firs (Abies) and false box (Gyminda)--not interested in obtaining. Sargent, however, was counting nomenclatural transfers like Carya rarely satiated, and he requested that Munson to Hicorius. In 1895, however, Sargent proreturn to collect half-a-dozen specimens of branches with winter buds.12 posed another fourteen names--many of them The following March, Sargent rode the rails oaks (Quercus)--suggesting he was becomto Texas to see the inexplicable hickory himing more confident of his own taxonomic eye. self. He also stopped in central Arkansas, where Those numbers continued to grow, and by he botanized in the alluvial bottomlands near 1907, he had added over three hundred names the town of Van Buren with the engineer of the in Crataegus alone. 40 Arnoldia 76\/1 ? August 2018 municipal waterworks, George M. Brown, who was an avocational student of the flora. Munson and Brown were new collaborators for Sargent, and he took fondly to both. When he returned to Brookline, hickory propagules had already arrived from Brown. Sargent requested flowering specimens from both men--apparently his trip had missed the spring flush--and although he reminded them to gather fruiting specimens in the fall, Sargent ultimately returned to observe the plants himself. 13, 14 He visited both men in early October and also rendezvoused to talk about hickories with his longtime collector Benjamin Franklin Bush, who ran a general store near Kansas City, Missouri, and who had already proven himself a keen botanical observer for Sargent's hawthorn research. While he was travelling that fall, Sargent collected seed for at least nine Arboretum hickory accessions. Only one plant from this collecting trip is still growing at the Arboretum today, representing our oldest accession of the nutmeg hickory (Carya myristiciformis, accession 6048*C), a rare species, which Sargent collected in Arkansas. It is now an impressively straight-trunked specimen in the center of the hickory collection, growing not far from a smaller-statured black hickory (C. texana, accession 12892*C), sent from Brown in 1912, and a pignut hickory (C. glabra var. megacarpa, accession 18062*A), which Bush collected in southern Illinois that same year. Sargent's enthusiasm was officially brimming. Hickory Problems If the unusual black hickory in Arkansas initially sparked Sargent's concerted investigation of the genus, publication projects breathed oxygen onto the flame. While Sargent began working on his first edition of the Manual of the Trees of North America, distilling his work on the fourteen-volume Silva into one comprehensive guidebook (published in 1905), he began simultaneously proposing and describing new taxa in serialized publications titled Trees and Shrubs: Illustrations of New or Little Known Ligneous Plants. These were released incrementally, and his research on hickories would appear in the final installment, published in 1913. As the publication date approached, he began firing off letters to collectors, urging them for information about hickories. Many of the correspondents were recent hawthorn collaborators--tried and tested in their ability to field ceaseless requests--although Sargent even turned to his old census agents, perhaps because their trees were already growing in the Arboretum collection. \"You used to be very keen about Hickories and I hope that you will have a relapse of the Hickory fever and make large collections again,\" he wrote to Letterman in 1911, even though the two hadn't corresponded significantly over the intervening years. \"The genus has got to be reworked and I am getting together as much material as possible for this purpose that it may make a better showing in the new edition of my Manual than it does in the first edition.\" 15 He also wrote to Ridgway the same year, and Ridgway responded with characteristically meticulous and detailed handwritten notes, and professed enthusiasm for the project. \"I have long been convinced that the genus is in sad need of overhauling,\" he wrote, \"and feel sure there are several more good species than are recognized in the books.\"16 When Sargent ultimately published his treatment on the genus in 1913, he proposed seven new species or hybrids along with an additional thirteen varieties--marking his first published effort to disentangle and redefine taxonomic parameters within the hickories. (One of these hybrids, Carya ? brownii, was based on an individual tree in the bottomlands of the Arkansas River, where it had puzzled Sargent and Brown back in 1909.) Yet this research on the hickories still proved unsatisfactory. Harbison--Sargent's faithful southeastern collector--had made extensive collections of hickories the same fall the report was published. \"I must say the more I see of them the more confused I become,\" Sargent wrote about material Harbison had sent from Georgia and Alabama, typing his frustrated missive on New Year's Eve. \"It is evident, I think, that we cannot depend much on the fruit as I once supposed we could and that we must try for other characters, bark, habit, location, habitat, winter-buds, pubescence, etc. I do not suppose that there are a great many species but the trouble is to limit what there Hickory Taxonomy 41 are. It seems to me that it will be impossible to properly know them without a great deal more field observation.\"17 Notes like this became a recurring refrain over the next several years, as he repeatedly asked Harbison, Palmer, and others for additional information about specimens that had arrived. Sargent would ultimately publish his final taxonomic treatment of the genus in 1918, when it appeared in the Botanical Gazette. He proposed more than twenty additional taxa, many of them varieties and forms. By the number of proposed names, this placed hickories in the top three genera that Sargent had studied, behind only oaks and, of course, hawthorns. Sargent closed that final report by describing thirteen individual trees that had been observed by John Dunbar, the assistant superintendent of the Parks Department in Rochester, New York. Sargent provided precise notes about the color of the branches and the shape of the fruit. None of these thirteen plants resulted in accessions that are still growing in the Arboretum collection, although we still have eight plants (representing seven unique wild provenances) from Dunbar and his collaborator Bernard H. Slavin. Sargent praised the collectors and noted that no region had been more \"carefully examined\" for hickories than western New York. To Sargent, the discovery of confounding individuals there simply proved that other regions needed to be studied with the same rigor. If so, he suspected additional taxa would be discovered. Nevertheless, hickories faded from Sargent's correspondence, and he would never publish another taxonomic treatment of the genus. Whether this report absolved what Sargent described as the \"hickory problem,\" however, remains unclear. Several months before the report was published, he wrote a letter to Reginald Somers Cocks, a professor at Tulane University, who had been a frequent correspondent on the genus. \"I have about finished up what I can do with Carya,\" he wrote, \"not a very satisfactory work.\"18 More than American Sargent based his description of Carya ? dunbarii on herbarium specimens from this tree, which John Dunbar had observed near Golah, New York. Richard Horsey, who worked with Dunbar at the Rochester Parks Department, photographed the tree and an unnamed man (could it be Dunbar himself?) in December 1918, shortly after publication of Sargent's final report. Notably, during much of this period, hickories were one of the few tree genera that appeared unique to the eastern North American flora. In the unattributed Garden and Forest article from 1889, the author--again, presumably articulating ideas approved by Sargent--had described the wood and fruits in superlative terms. \"As a nation we owe much to the Hickory tree, and we have good and just reason to be proud of it,\" the author wrote, even suggesting that the lightweight yet durable carriages crafted from hickory had allowed equestrian breeders to develop the American trotting horse, \"that race of horses which every American looks upon in his heart of hears with joy and admiration.\" 42 Arnoldia 76\/1 ? August 2018 Then, in 1915, Sargent received herbarium specimens of a Chinese hickory from the collector Frank Nicholas Meyer, who had first observed the fruits being sold at a market in Hangzhou, in eastern China. Sargent acknowledged receiving the specimens in a letter to Meyer the following January, and, of course, he requested more information about the size and abundance of the trees, not to mention photographs.19 Sargent's intrigue about the discovery, however, is perhaps most evident in his account of the species in Plantae Wilsonianae. Sargent edited the three volumes, published between 1913 and 1917, yet of nearly eight hundred names proposed for new Chinese plant taxa (not counting nomenclatural transfers), most came from other Arboretum staff--prominently Alfred Rehder and Ernest Henry Wilson--as well as European colleagues like Camillo Schneider and Bernhard Koehne. Sargent authored only seven new names: six hawthorns and one hickory--what he called Carya cathayensis. \"Since the finding in China of a species of Liriodendron [tulip tree] and of Sassafras, previously believed to be monotypic genera of eastern North America, no addition to our knowledge of the distribution of the trees of the northern hemisphere is so important and interesting as Mr. Meyer's discovery of a representative of the genus Carya in Asia,\" Sargent declared in the publication, noting that progressively few genera appeared unique to eastern North America. \"In China,\" he continued, using a tone that could suggest a trace of disappointment, \"there are many endemic trees.\"20 Strangely, Sargent never acquired seed of this species from Meyer-- perhaps suspecting they would be unable to grow at the Arboretum, given that it was discovered in the humid subtropics--and the only material ever collected (by Peter Del Tredici in 1989) never made it out of the greenhouse. It is currently on the list of desiderata for the Campaign for Living Collections. In 1915, Sargent learned that the American hickories like the charismatic shagbark (Carya ovata), photographed in Rochester, New York, above, had Asian relatives. That summer, Frank Meyer photographed a large Chinese hickory (Carya cathayensis) on the edge of a bamboo grove in Zhejiang Province. \"The wood is said to be tough and strong and appreciated as handles for agricultural tools,\" Meyer noted. Hickory Taxonomy 43 Confidence in the Commonplace Botanists would eventually begin consolidating many of the hickories Sargent had named. Because so many of the taxa were varieties, many of the names have been dropped in recognition of more morphologically diverse conceptions of each species. \"Phenotypic variation from tree to tree is often considerable and difficult to quantify,\" Donald Stone wrote in his treatment of the American species in the Flora of North America, published in 1997. \"Most of this variation undoubtedly results from adaptation to local and regional conditions; hybridization has probably played a subtle role as well.\"21 Stone included eleven species--down from the fifteen in Sargent's final report--and referenced another seven species globally (most in eastern Asia, one in Mexico), although these numbers have fostered ongoing debate.22 In the spring of 1918, as Sargent was wrapping up work on his final hickory report, he wrote to Harbison, suggesting that certain hickory species had been neglected by botanists, given their general abundance and familiarity.23 It was this fundamental spirit that inspired what must be, even still, one of the most widespread and detailed morphological studies of the North American members of the genus--an impossible endeavor without the nuanced observations by Sargent's cadre of mail correspondents. This collaborative effort also provided the centenarian core for the Arboretum's robust collection of hickories, which was one of our first to be accredited by the American Public Gardens Association's Plant Collections Network in 2002 (back when the network was known as the North American Plant Collections Consortium). Current field-collecting efforts continue to prioritize the genus, maintaining the intellectual passion of Sargent and his collaborators long ago: the confidence that even among the \"plants which have been considered too common to collect,\" something new can always be discovered. Endnotes 1 Sargent, C.S. 1914. Sargent to T.G. Harbison, 2 March 1914 (volume 8, page 8). Charles Sprague Sargent (1841? 1927) papers, Arnold Arboretum Horticultural Library, Harvard University. (All Sargent correspondence below from the same archive, unless otherwise noted.) 2Sargent to E.J. Palmer, January 19, 1915 (volume 8, page 308). 3For the term \"special agent\" applied for collectors other than Sargent himself, see Sargent, C.S. 1902. Silva of North America, 13 (pp. 79?80.) 4Ridgway, R. 1882. Notes on the native trees of the lower Wabash and White River valleys, in Illinois and Indiana. Proceedings of the United States National Museum, 82(1): 49?88. 5Letterman. G.W. 1881. Letterman to G. Engelmann, April 12, 1881. Engelmann papers, Missouri Botanical Garden. 6Engelmann, G. 1881. Engelmann to G.W. Letterman, April 14, 1881. Engelmann papers, Missouri Botanical Garden. 7Sargent, C.S. 1902. Silva of North America, 13 (pp. 79?80). Boston and New York: Houghton Mifflin and Company. 8 Sargent, C.S. 1889. Notes upon some North American trees--XI. Garden and Forest, 2(83): 459?460. 9Anon. 1889. The shell-bark hickory. Garden and Forest, 2(83): 460?461. 10Sargent to R.T. Morris, October 21, 1908 (volume 6, page 161). 11Sargent to T.V. Munson, October 19, 1908 (volume 6, page 160). 12 Sargent to T.V. Munson, November 7, 1908 (volume 6, page 190). 13Sargent to G.M. Brown, March 31, 1909 ( volume 6, page 383). 14Sargent to T.V. Munson, March 31, 1909 (volume 6, page 384). 15Sargent to G.W. Letterman, June 7, 1911 (volume 7, page 122). 16Ridgway to C.S. Sargent, August 6, 1912. 17 Sargent to T.G. Harbison, December 31, 1913 (volume 7, page 908). 18Sargent to R.S. Cocks, June 19, 1918 (volume 9, page 296). 19Sargent to F.N. Meyer, January 17, 1916 (volume 8, page 618) . 20Sargent, C.S. (Ed.). 1917. Plantae Wilsonianae, 3. Cambridge: The University Press. 21Stone, D.E. 1997. Carya. Flora of North America North of Mexico, 3. New York and Oxford: Flora of North America Association. 22 Zhang, J.B., et al. 2013. Integrated fossil and molecular data reveal the biogeographic diversification of the eastern North American disjunct hickory genus. PLOS ONE, 8(7): 1?13. 23Sargent to Harbison, June 19, 1918 (volume 9, page 294). Jonathan Damery is Associate Editor of Arnoldia. "},{"has_event_date":0,"type":"arnoldia","title":"Existing through Change: Quercus alba","article_sequence":4,"start_page":44,"end_page":45,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25645","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15e896d.jpg","volume":76,"issue_number":1,"year":2018,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Existing through Change: Quercus alba Michael S. Dosmann O n a magnificent bird's-eye-view map of the Arnold Arboretum, prepared in 1927, paths and hillsides, collections and trees are depicted in such clear detail that you can easily imagine gazing down from a floating hot-air balloon (see pages 18?19). The map hangs in the Arboretum library, and perhaps the most striking specimen is a tree drawn on the northeast slope of Peters Hill, a lone shepherd guarding an endless flock of Charles Sargent's beloved hawthorns (Crataegus). This venerable singleton is a double-leadered white oak (Quercus alba), now towering above the Arboretum's crabapple (Malus) collection, which replaced most of the hawthorns after World War II. The tree reaches an impressive spread of 28 meters (92 feet) and height of 17.2 meters (56 feet)--the southwestern bole is slightly taller--but most notable is its girth, no doubt exaggerated by the twin stems, which are swollen with crown gall (Agrobacterium tumefaciens). A slight seam meanders through this burly base, suggesting the stems are fused at ground level, where the diameter is 2 meters (6.5 feet). Impressive is an understatement. Despite its size (and obvious age), a glance at the tag reveals the tree's accession number: 3462010*A, meaning it was accessioned just eight years ago. Ordinarily, the Arboretum receives accessions as wild-collected seed, nursery purchases, cuttings from sister gardens, or other means (there are thirty-three different options), yet this tree is an \"existing plant.\" It was officially given its unique accession number after it had been found growing in place. Of the 14,722 accessioned plants in the permanent collections, almost 20 percent are existing plants. Over half of these were accessioned since 2007, through an initiative led by Manager of Plant Records Kyle Port, which officially brings important specimens from managed areas into our systems to be measured, tracked, and studied. Even without an accession number, these trees with opaque provenance received arboricultural care; curatorial recordkeeping adds collections value. Our oak first shows up in Arboretum archives on a detailed 1894 topographical survey of Peters Hill made for the City of Boston, a year before this area became part of the Arboretum. At first glance, the tree appears to be marked with an 8, but this figure actually represents the two stems. To establish a firmer age, I recently extracted an increment core and counted the rings from the southwest stem, 138 centimeters (54 inches) above the ground. The wood was hard and intact most of the way until I approached the center and hit a soft pocket. Even so, with 70 percent of the core intact, I counted 142 rings. Assuming the unrecovered portion represented an additional 60 rings, that leader reached its position in the 1810s; germination would have been several years earlier. With this information, we've affirmed the tree to be of local, wild provenance. In 1937, Hugh Raup shared his thoughts about several existing white oaks on Bussey Hill. One tree's rings, counted after a fatal lightning strike in 1931, dated to 1666. Raup pondered what this tree had witnessed, particularly landscape changes from forest to agriculture to woodland again. Among all the changes, however, Raup stated \"the later scenes [of the Arboretum] are the strangest.\" The bizarre combination of exotic and local plants surpassed the previous centuries' revolution: the \"great white wings\" of the dove tree (Davidia involucrata) and \"impossible maples with copper-colored bark that peels off in thin sheets\" (Acer griseum). Change is inevitable, and the Arboretum's naturalistic landscape is no exception. In another century, perhaps a new collection will replace the Peters Hill crabapples just as they eclipsed the hawthorns. With good graces, I hope our white oak is around to bear witness to the transformation, for as Raup wrote of the Arboretum, \"the only continuity is in the inherent charm of the place and in the lives of the ancient oaks.\" Literature cited Raup, H.M. 1937. The Chinese Collection. Bulletin of Popular Information, Series 4, 5(5): 25?28. Michael S. Dosmann is the Keeper of the Living Collections at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23462","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14eb728.jpg","title":"2018-76-1","volume":76,"issue_number":1,"year":2018,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"What the Rose Knows: Navigating Rosaceae at the BRC","article_sequence":1,"start_page":2,"end_page":16,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25644","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15e8928.jpg","volume":75,"issue_number":4,"year":2018,"series":null,"season":null,"authors":"Bowman, Erica","article_content":"What the Rose Knows: Navigating Rosaceae at the BRC Erica Bowman T When we develop botanical collections, plant specimens are the stars, and the genius lies in their artful and memorable arrangement. Strict taxonomic accuracy has never been our primary role. When the Julie Moir Messervy Design Studio (JMMDS) was hired to help redesign the Arnold Arboretum's Bradley Rosaceous Collection (BRC) in 2007, the objective was simple. We wanted to improve upon the sense of space in the existing garden without disturbing the living collection or dishonoring its legacy. We wanted the garden to be lightly educational, a place where visitors could experience taxonomic order in a beautiful, satisfying setting. We were determined to use an organizational MENG LI aken literally, Gertrude Stein's famous phrase \"Rose is a rose is a rose\" might have raised the collective eyebrows of a few scholars of the rose family (Rosaceae). After all, more than four thousand species and one hundred genera make up this complex family of woody and herbaceous plants. Robert Frost rendered these botanical relationships more accurately, writing, \"The apple's a rose, \/ And the pear is, and so's \/ The plum, I suppose,\" yet even he failed to mention less-lyrical genera like Spiraea, Cotoneaster, or the elusive Prinsepia, not that anyone could blame him. Frost was no more a taxonomist than Stein, or, ahem, a lot of landscape designers. Like poets, we landscape designers are primarily artists. Gertrude Stein's famous line \"Rose is a rose is a rose\" first appeared in the 1913 poem \"Sacred Emily.\" Robert Frost co-opted part of the phrase to open \"The Rose Family,\" a poem published in 1928. This specimen of Macdougal's rose (Rosa nutkana ssp. macdougalii, 516-78*MASS) was wild-collected in British Columbia. DAVID BASLER AND MARGARET KOSMALA, HARVARD UNIVERSITY Bradley Rosaceous Collection 3 Swooping bed lines in the BRC demonstrate Olmstedian sensibilities. In this photograph, taken above the east beds in May 2017, elements of the original 1980s layout can be detected as swaths of darker turf. The main flowering specimens shown in each bed, clockwise from the bottom left, are Wilson spirea (Spiraea wilsonii, 545-93), Henry spirea (S. henryi, 1121-86), another Wilson spirea (953-85), and smooth oriental photinia (Photinia villosa, 934-85). Dawson Pond brims with spring rain on the left. structure that visitors could intuitively understand through passive experience, whether or not they were botanical scholars. Mostly, we hoped that the garden would appear rejuvenated, while still reverent to the surrounding landscape that was developed by Frederick Law Olmsted and Charles Sprague Sargent, the first director of the Arboretum. All of this--simple as it initially seemed-- was no minor responsibility. The list of stakeholders was vast and intimidating. There were donors to consider and staff to please. We had to be respectful of the Arboretum's legacy, honoring the institutional position within both Harvard University and the City of Boston. We even had to consider the reputation of the esteemed plants themselves. Julie Moir Messervy had already been inducted into the Arnold's folds. In addition to lecturing there on numerous occasions, she designed the Linda J. Davison Memorial Path that winds along Bussey Brook. She also planned the landscape surrounding the granite bench that overlooks the BRC. Despite my role as project manager, I was a relative newbie on this scene, having just started working with JMMDS. I completed my MLA at Cornell in 2003, where, under Professor Don Rakow, I studied some of the world's most hallowed gardens: Longwood, Kew, and the Arnold Arboretum. My arrival at the BRC felt monumental to say the least. I knew I was on sacred grounds. ARNOLD ARBORETUM ARCHIVES 4 Arnoldia 75\/4 ? May 2018 The location now occupied by the BRC was formerly known as the Shrub Collection, a series of long, straight rows that existed from 1885 through about 1982. By the time the Arboretum was first systematically mapped in the late 1930s, roses--that is, specimens of Rosa--occupied the center of the Shrub Collection, the most dominant genus growing there. When we began our work, the BRC contained over forty Rosaceous genera and up to four hundred different taxa. Keeper of the Living Collections Michael Dosmann (who then held the title of Curator) provided a \"desiderata\" of seventy-five additional taxa that the Arboretum wanted to incorporate into the design. The existing collection was not presented in an obvious order within the family itself. According to feedback from gardeners and staff, many visitors didn't understand the taxonomic connection between woody Rosaceous genera and plants in the Arboretum at large. Some plants were duplicated; others were overgrown. And visitors and staff both complained that most of the Rosas had a very short season of bloom, primarily in the month of June, and that even then, many of their differences were subtle and hardly obvious. Much of the time, the roses presented themselves as a sea of thorny stems. Each rose in the vast collection could be mistaken as just another rose. A rose was a rose was a rose. Moreover, although the beds were clearly defined, we wanted to improve upon the overall sense of place and intensify the experience of journey. We use \"sense of place\" to mean everything that defines a unique and recognizable landscape. Take Central Park, for example: A visitor can look up at the skyscrapers and then look down to see the Reservoir, the Great Lawn, and the stony bridges. Even with eyes closed, they will experience the sounds and smells that define the iconic city park. All of Bradley Rosaceous Collection 5 this contributes to their sense of being located in a memorable place in the world--exactly what we wanted for the BRC. ARNOLD ARBORETUM ARCHIVES Our creative process always involves iteration, beginning with the development of an overarching \"big idea.\" We considered a wide range of thematic concepts for the BRC. We envisioned the botanical structure of a rose flower, for example, with its five petals and sepals as a template for bed shape and placement. We compared contemporary and classic interpretations of the rose in folklore, religion, literature, and romance. These ideas were manifested in overlays of pencil upon tracing paper, often times starting and ending as scrawled words in the margins: \"What Rosaceous plants are mentioned in Shakespearean plays? What was the significance of the rose in world religions? How are Rosaceous plants used medicinally?\" We sketched routes and journeys for the pedestrian to travel from place to place, borrowing inspiration from Olmsted's pastoral style. We sketched potential bed outlines, envisioning new outdoor rooms and meeting spaces, new places to sit, and new views to see. The resulting documents were beautiful but chaotic, a palimpsest of possibilities. Still, the big idea hadn't taken hold. At JMMDS, we recognize that a big idea can evolve from many places. When Julie worked with Yo-Yo-Ma on her design for the Toronto Music Garden, they used Bach's Suite no. 1 in G Major for Unaccompanied Cello as a musi- The BRC was endowed by Eleanor Cabot Bradley, pictured in white at the dedication ceremony on June 12, 1985. Her daughter Elizabeth Cabot Sluder is shown planting a ceremonial rose. Arboretum Director Peter Ashton stands second from the right, beside Horticulturist Gary Koller, who crafted the original garden design. In 2007, Sluder graciously supported the garden renovation. JULIE MOIR MESSERVY DESIGN STUDIO 6 Arnoldia 75\/4 ? May 2018 This 2008 renovation map uses color codes to indicate which plants were included in each of the old Rosaceous subfamilies. Rosoideae is shown in red, Maloideae in blue, and Spiroideae in green. (This particular bed had no members of Prunoideae.) Marginal comments describe horticultural characteristics and backstory for specific accessions. cal template. The resulting garden is a celebration in six movements--prelude, allemande, courante, sarabande, minuet, and gigue--all rhythmically expressed as wooded paths, great spiraling landforms, bee-filled plantings, and oversized grass steps. JMMDS's design for the Inspiration Garden at the Shore Country Day School in Beverly, Massachusetts, on the other hand, took the feel and form of the school's mascot, the beaver. We recreated habitat with native plantings and erected an outdoor classroom shaped like a beaver lodge. While brainstorming the big idea for the BRC, we looked again at a base map the Arnold had provided. It included a key with four subfamilies: Rosoideae, Spiroideae, Maloideae, and Prunoideae. As it turned out, these subfamilies had already become outdated, yet from a design perspective, the groupings created a useful ethnobotanical order. Rosoideae traditionally included, among other things, roses and brambles (Rosa and Rubus), so for that subfamily, we imagined a classic rose garden with arbors, benches, and other ornamental features. Maloideae contained apples (Malus) and hawthorns (Crataegus), along with a number of larger tree species, so we thought of an orchard planting. Prunoideae inspired us to think of ornamental trees blooming at a Japanese sakura festival, because the most dominant genus in the group includes cherries and plums (Prunus). Spiroideae--with members like nine-bark (Physocarpus), pearl-bush (Exochorda), and spirea (Spiraea)--brought to mind a well-ordered shrub collection. Obsolete or not, this historical classification helped us organize a series of places, each with their own beauty and meaning, and all connected by a journey through the larger family. We tentatively named these components the Bradley Rosaceous Collection 7 that would allow for a sort of compromise of ideals between taxonomy and design. \"Let's call it a `collection of arborescence' rather than a literal orchard,\" one person offered. \"Fruiting arborescence?\" we countered. To that they agreed. By \"arborescence,\" of course, we meant \"trees.\" That is the way botanists and landscape designers talk. Sometimes we pontificate in French, Latin, and Greek, other times in ultra-syllabic versions of regular English (\"fenestration\" instead of \"windows,\" \"rectilinear\" instead of \"rectangular\"). Yet, to communicate with the rest of the normally speaking world, we needed to tone down our jargon. So we retracted the place names KYLE PORT Rosoideae Roundabout, the Maloideae Meander, the Prunus Promenade, and the Spiroideae Stroll. Keeping this old taxonomic organization had the added benefit of allowing us to maintain the larger, less-movable trees in their place, given that many were grouped taxonomically already. This allowed the greatest amount of transplanting and bed realignment to happen with the more manageable roses and shrubs. It also made the plant groupings easy to comprehend on a basic level. We hoped that a visitor might be able to travel from area to area and immediately sense order within the groupings. The Arnold staff and donors tried to help us make this plan work. They steered us away from the literal interpretation of the old subfamilies and suggested slight verbiage shifts JMMDS imagined a section of flowering cherries that would invoke the spirit of hanami, the Japanese tradition of celebrating the transient beauty of spring. Here, visitors gather in the Prunus Promenade on April 30, 2013. 8 Arnoldia 75\/4 ? May 2018 MENG LI Maloideae Meander and Spiroideae Stroll, which were a mouthful anyway, but we maintained the idea of shruband orchard-like collections. Rosoideae Roundabout seemed to work, especially since this subfamily was still legitimate, but the name was reduced to the Rose Roundabout for good measure. The Prunus Promenade was too lyrical to abandon. We had found our big idea. MICHAEL DOSMANN Cost, availability, and establishment time almost always influence plant selection and expectations. When we design a garden, we make plant choices based on nursery catalogs and availability lists. We're also used to waiting a bit before our designed landscapes grow into themselves. That is just part of the practice. For a perennial garden to reach its prime, three years is the normal window. Most practitioners plan for a fifteenyear grow-in period for trees. These establishment times assume that, at installation, perennials are gallonsized and trees have a 1.5- or 2-inch caliper trunk. Even this can be too small and slow for some clients. In those cases, we can crane in mature twenty-foot trees with six-foot root balls for an immediate impact. This is costly but instantly satisfying. The BRC is no average garden, however, and the Arnold certainly doesn't require instant satisfaction. As part of the larger living collec- Provenance is important for plants in the BRC, whether a specimen of tions, every plant in the BRC has a Japanese kerria (Kerria japonica, 237-2008*B), which was wild-collected in Gunma Province, Japan, or a `Donald Wyman' crabapple, propagated from standardized accession number, and the original specimen on Peters Hill (23254*A). all new acquisitions must meet the requirements laid out in the Living Collections these considerations in mind, it made sense to Policy. Many new accessions are wild-collected tackle the Rose Roundabout first, because those as seed and then grown in the Arboretum's nursplants were the quickest growing and most ery. Even the addition of new cultivars can take easily movable. The completion of the Prunus time. For instance, the crabapple Malus `Donald Promenade and orchard collection could wait. Wyman,' which the Arboretum introduced in 1970, was repropagated from the original tree, New and unfamiliar plant material was another even though a large specimen could have been obstacle for our team to clear. Few genera in purchased from a commercial nursery. With the BRC were completely new to us, but more MENG LI often than not, we were in the practice of working with more widely available, commercially propagated plants. We have had much less experience dealing with straight (and often obscure) species like the ones that fill the BRC. Delving into Rosoideae was particularly demanding with a large number of plants that seemed, at first glance, to have similar outward characteristics in the off-season: almost all were thorny and shrubby. We needed to learn more about the specimens in order to make design decisions. As the resident plant person at JMMDS, I was sent to the Arnold to receive some additional schooling. I spent days walking around with Kit Ganshaw, the long-time BRC horticulturist, discussing the particulars of each plant. Within Rosoideae alone, I learned that the Arnold had a large planting of Rosa `Justine Mee Liff', a cultivar named for the first woman to serve as Boston parks commissioner. The cultivar `Duchesse de Montebello' was an important gift from the Bradley family and needed to be preserved, especially as we ultimately reduced the garden in that location to make way for a new path. While the yellowflowered incense rose (R. primula) is the first rose to bloom in the garden, the single specimen of climbing prairie rose (R. setigera) was significant in that it blooms in July, after Other plants with known wild origins include this Henry spirea (Spiraea most other species are done. The henryi, 302-84*A), which was repropagated from an accession Ernest Henry Wilson collected in western Hubei, China, and this specimen of Rosa zalana apple-scented blossoms of the sweet- (928-78*G), which was collected north of Budapest, Hungary. brier rose (R. eglanteria) feature in the writings of Shakespeare. The whitedeemed a very special plant because it came stemmed bramble (Rubus cockburnianus) profrom the Duke of Buccleuch's estate garden-- vides great winter interest with its showy stems, considered one of the premier rose collections and Sweginzow's rose (Rosa sweginzowii) is when Sargent received a shipment of cultivars noticed for its glorious hips. Mount Omei rose in 1914--and it may be the only specimen of (R. omeiensis) grows up to twelve feet tall, has `Plato' remaining in cultivation. The claim of bright red thorns and an atypical arrangement fame for serafin rose (R. serafinii), according to of four petals to its flower. A trellis-worthy culKit, is that is has the \"nastiest hooks\" in the tivar of Scotch rose (R. spinosissima `Plato') was whole collection. Whew! MENG LI Bradley Rosaceous Collection 9 10 Arnoldia 75\/4 ? May 2018 Rose by Rose, Cobble by Cobble: A Garden Renovation Michael S. Dosmann and Stephen Schneider I t was January of 2007, Michael's first week as Curator of Living Collections, when former Arboretum Director Bob Cook presented three immediate curatorial priorities: reenvision the Arboretum's Living Collections Policy, implement environmental monitoring systems (including GIS), and \"do something about the rose garden.\" Despite several decades of care and attention since its intentional design in the early 1980s, the Bradley Rosaceous Collection (BRC) had become overgrown and lost its focus. Both the addition and subtraction of plant material had been limited. From a curatorial perspective, the dense plantings were a nightmare to label and keep authentic. It was time to make things right for a family with deep ties to the Arboretum by holding ourselves accountable to high levels of care and curation. In crafting a new vision for the BRC, we wanted to celebrate plant diversity writ large: not just botanical diversity provided by wild-collected woody species, but also cultivated diversity so richly illustrated by old and new cultivars. Accessions in the BRC, like those elsewhere in the Arboretum, would be valued based upon their full documentation and provenance, not simply a hierarchy of wild over cultivated origin. The attention to cultivated diversity would also solve a problem for us: come midsummer, few things in the garden bloomed. By adding new cultivars of roses (and other species), we could increase the garden's display potential. Around this time, we were dramatically rethinking how we deployed resources for horticulture. Under the leadership of former Deputy Director Richard Schulhof, the Arboretum launched an initial Landscape Management Plan (LMP) in 2007. The LMP prescribed expectations for arboricultural and horticultural care throughout the collections, including the BRC. Simultaneously, we undertook a curatorial review of all plants in the BRC to determine which lineages to preserve. The garden was packed, and the last thing we wanted was to renovate again in ten years. Even so, we still lacked the perspective that only a garden designer could provide. We needed innovative ways to organize plants that would maximize display potential (including for \"BIO\" plants that possess \"botanical interest only\"). We also needed to improve the visitor experience. Without a formal entrance, the garden lacked a sense of arrival. Circulation also demanded attention. While the existing bed configuration (full of tall, dense shrub masses) created intimate garden rooms, it also inhibited visitor exploration due to fear of the unknown or even concerns for safety. In Julie Moir Messervy Design Studio, we found technical expertise to help us tackle these and other problems. Erica and Julie provided both creative genius and sensitivity to the project (and its idiosyncratic client). Although we didn't intend to completely redo the five-acre site, we were seeking a major renovation. Shortly after the project launched in 2008, the Great Recession hit, forcing us to adjust the initial timeline that had called for a single season of renovation using contracted labor. The budget also caused us to rethink aspirations for formal paths and benches, at least initially. Instead, we used our in-house team of horticulturists and interns for the renovation, extending the bulk of the project from one year to three. With JMMDS's new plans in hand, we completely deconstructed some beds, while others were reshaped or constructed anew. This involved handling some 10,000 double cobbles, and a parade of pallets formed along Forest Hills Road. Shifting stones. Double cobbles were installed around a new bed in September 2010, while exposed soil remained visible in the footprint of an old bed (at right). The new plans identified plants to remove, as well as plants that could shift to new locations in the BRC. In some cases, the move could occur instantly, but because of the renovation's phased approach, other plants were transplanted to a temporary location near the Dana Greenhouse and Nursery's south nursery until new beds could be created. This intricate dance kept staff on their toes as they ensured plants were labelled and documented at all times. The greenhouse and nursery staff maintained these, while repropping important accessions from the BRC and beyond. Before we knew it, old beds morphed into turf and new beds appeared. Fresh rose cultivars extended the bloom season in the Rose Roundabout as well as other spaces in the garden. The Prunus Promenade materialized as trees were planted out (including additional cultivars). By the spring of 2011, most of the work was completed, including the installation of Peter Andruchow's beautiful arbor in the Roundabout. This destination for climbing roses was dedicated to Elizabeth Cabot Sluder (daughter of Eleanor Cabot Bradley) during an event with family and staff on June 18. We continue to implement elements of the JMMDS plan for the Bradley Rosaceous Collection. The orchard, comprising a diversity of harder-to-acquire germplasm, has taken some time to fill in, but as of this spring, we are about three-quarters of the way there. In 2013, we hired Peter to construct a second arbor, identical to the one in the Roundabout, which serves as an entry portal between two new beds along Meadow Road. We have also installed granite seating--akin to the bench Julie helped design above the BRC years before--as part of the Arboretum's new Commemorative Bench Program. Gardens, ever changing and dynamic, evolve and grow. The nearly 150-year history of this location, near the Forest Hills Gate, is a perfect example. Michael S. Dosmann is Keeper of the Living Collections and Stephen Schneider is Director of Operations at the Arnold Arboretum. MICHAEL DOSMANN MICHAEL DOSMANN Bradley Rosaceous Collection 11 ARNOLD ARBORETUM GIS\/HARVARD PLANNING AND PROJECT MANAGEMENT 12 Arnoldia 75\/4 ? May 2018 This 2015 photograph shows the BRC as redesigned by JMMDS, while the yellow outlines depict contours of the original 1980s design. The Rose Roundabout is located on the left, while the beds to the right primarily showcase Rosaceous shrubs. Notice that cherries are flowering pink and white in the Prunus Promenade. The orchard collection is located in the lawn near Willow Path, which runs along the top side of the collection. Additionally, we identified problem plantings. Some masses like prickly rose (Rosa nipponensis) needed reduction. Others, like wood rose (R. gymnocarpa) needed increasing. R. ? nitidula was too tall for the front of the garden, while interior rose (R. woodsii) and an unusual Taiwanese species (R. transmorrisonensis) were too short for the back. Species like red-spined rose (R. nitida) and swamp rose (R. palustris), both wet-loving, might have been better suited near the ponds, and longtooth rose (R. longicuspis) needed a trellis. Some of the catalogued plants were even missing from the field, including the namesake Arnold rose (R. ? arnoldiana `Arnold'). Clearly, reorganization was in store. Ultimately, we would suggest removal of sick and duplicated plants, reduction of colonies, and organization by height, habit, and bloom period to create densely alluring focal points throughout the summer. We fine-tuned the shape of the beds to improve circulation, pauses, and views. We also designed a main entrance and a circumferential route around the garden. Still, even with restructuring, we were concerned that some distinctions between plants might be too subtle for the average visitor. Our instincts wanted to make the whole collection more photogenically engaging. We wanted to intersperse herbaceous rose family JULIE MOIR MESSERVY DESIGN STUDIO Bradley Rosaceous Collection 13 KYLE PORT This illustration shows the initial design concept for the entrance arbors. plants, like lady's mantle (Alchemilla mollis), meadow sweet (Filipendula rubra), and avens (Geum spp.), between colorful rose hybrids. The Arnold pushed back on that idea, favoring a more dignified, uncomplicated (if not staid) order to things. Our mixed-garden proposal also complicated the maintenance practice at the time, which emphasized a clean contrast between mulch and woody specimens. That approach is easier to maintain, since any invading plant can easily be identified as a weed. Our compromise was to integrate longer-blooming hybrid roses like `Radtko' (Double Knockout?), `Chuckles', `Bucbi' (Carefree BeautyTM), and `AUSbord' (Gertrude Jekyll). We still dream of incorporating more herbaceous members of the rose family. In the very least, we want to eliminate views of mulch. To honor philanthropist Elizabeth Cabot Sluder, we designed a new arched entrance to the Rose Roundabout and put metal artist Peter Andruchow of Wovensteel to work. JMMDS designer Jana Bryan incorporated features from the Arboretum's wrought-iron main entrance into the design, and she also added hallmark puddingstone boulders at its base. We selected a delicate, pink climbing rose (Rosa `New Dawn') to clamber across the archway, a departure from the shrubbier citizenry of the Roundabout. The plants were shielded within the protective hollow of the arch's tri-scrolled feet. Rosa `New Dawn' (712-2010*A) grows over the Rose Roundabout entrance. When we evaluated existing plants in other parts of the garden--the orchard, the Prunus Promenade, and the shrub collection--it was mostly for location, health, quantity, and appearance. Unlike the Rosas, few of these plants came with noteworthy S. A. PFEIFFER 14 Arnoldia 75\/4 ? May 2018 The BRC redesign includes two wrought-iron arbors, one at the entrance to the Rose Roundabout, the other, pictured here, along Meadow Road. backstories, other than a rare Chinese species of mountain ash, Sorbus yuana, which was one of the first accessioned in the United States. We identified some plants as stand-out ornamental specimens, including Toringo crabapple (Malus sieboldii), intermediate shadbush (Amelanchier intermedia), and petalless cherry (Prunus apetala), along with the grove of quince trees (Cydonia oblonga). We noted that the trio of Siberian crabapples (Malus baccata) were beautiful trees that could offer a transition between the orchard and shrub collections. We also catalogued a large and beautiful cultivar of black cherry (Prunus serotina `Cartilaginea'), a tall and stately example of Gray's chokecherry (P. grayana), and an early blooming favorite of visitors, cyclamen cherry (P. cyclamina). One unconventional hybrid, (? Sorbaronia sp.) ? (? Sorbocotoneaster sp.), was so appealing that Julie marked it as something we should find in the trade. We haven't seen it yet. In order to create distinct areas within these portions of the landscape, we needed to move a lot of plants around. One of our aims was to remove mulch beds around large standard trees, allowing the trees to spring directly from the lawn, while reserving beds for shrub groupings. Additionally, we wanted to improve upon the beauty and understandability of the space by reducing clutter, redundancy, and confusion. This meant moving plants like Nantucket shadbush (Amelanchier nantucketensis) out of the Prunus Promenade, despite their beauty. To give more space to the handsomely fruited medlar (Mespilus germanica), it was necessary to remove a Wilson spirea (Spiraea wilsonii) MENG LI Bradley Rosaceous Collection 15 KYLE PORT This highland drophip rose (Rosa oxyodon, 295-2008*A) was wild-collected on the north side of Lake Sevan, Armenia. that had invaded its folds. This was fine by Kit, who was very comfortable reducing size and clutter through pruning. It was actually Kit who introduced me to the term rejuvenation pruning, the process of pruning a shrubby plant to the ground in order to bring about a new and improved flush. While I was not in the practice of making such sudden and brutal cuts on slower-growing plants, the Arnold, on the other hand, is a practical institution and nothing if not patient. This willingness to wait and withstand the awkward stages actually freed up decision making. Rather than removing a rangy or overgrown plant, we could opt to rejuvenate it instead. Specimens of Laland firethorn (Pyracantha coccinea `Lalandei'), flowering quince (Chaenomeles), and spirea (Spiraea) were all candidates for this treatment. Some plants were outright deaccessioned because of age, disease, redundancy, bad form, or incorrect placement. While a one-hundredyear-old dwarf Siberian peashrub (Caragana arborescens `Nana') escaped the chopping block, several large and lovely Kousa dogwoods (Cornus kousa) did not. Their removal from along Willow Path was perhaps the most painful part of the demolition phase for me. Although their execution papers were signed in the effort to purge non-Rosaceous plants, I mourn their loss to this day. Double Knockout? (Rosa `Radtko', 709-2010*A) was among the showy cultivars that extended the bloom season. It was photographed here on October 22, 2013. Renewal of the BRC opened spaces for planting new specimens. Since we were not in the position to know the exact plants for future accessions, we designed plantings by genus and habit (e.g., \"Malus, arborescent species.\") This allowed Arnold staff the freedom to grow and add new MENG LI 16 Arnoldia 75\/4 ? May 2018 Cyclamen cherry (P. cyclamina, 706-31*B) is native to south-central China. specimens as we moved into the future. In this respect, our role at the Arboretum has been different from other design jobs: We were not the most qualified plant experts on the team. We had to cede that role. It may be many more years before all of the plants have grown to maturity and our vision of the new BRC is realized. By then, scholarship surrounding plant classification will have continued to shift, as will the garden. This is the challenge faced by gardeners and taxonomists alike: constant momentum. The art of a garden is never static. It sprouts weeds and seeds, and begins to seek its own expression. This is why most garden designers like to remain involved in order to see their plans to fruition. Otherwise we are at the whim of nature, or to whomever holds the pruning shears. Erica Bowman is a senior landscape architect for Julie Moir Messervy Design Studio, where she manages projects in New England and beyond. She holds an MLA from Cornell University. She also writes for Horticulture Magazine. "},{"has_event_date":0,"type":"arnoldia","title":"Rose by Rose, Cobble by Cobble: A Garden Renovation","article_sequence":2,"start_page":10,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25641","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15e8126.jpg","volume":75,"issue_number":4,"year":2018,"series":null,"season":null,"authors":"Schneider, Stephen; Dosmann, Michael S.","article_content":"10 Arnoldia 75\/4 ? May 2018 Rose by Rose, Cobble by Cobble: A Garden Renovation Michael S. Dosmann and Stephen Schneider I t was January of 2007, Michael's first week as Curator of Living Collections, when former Arboretum Director Bob Cook presented three immediate curatorial priorities: reenvision the Arboretum's Living Collections Policy, implement environmental monitoring systems (including GIS), and \"do something about the rose garden.\" Despite several decades of care and attention since its intentional design in the early 1980s, the Bradley Rosaceous Collection (BRC) had become overgrown and lost its focus. Both the addition and subtraction of plant material had been limited. From a curatorial perspective, the dense plantings were a nightmare to label and keep authentic. It was time to make things right for a family with deep ties to the Arboretum by holding ourselves accountable to high levels of care and curation. In crafting a new vision for the BRC, we wanted to celebrate plant diversity writ large: not just botanical diversity provided by wild-collected woody species, but also cultivated diversity so richly illustrated by old and new cultivars. Accessions in the BRC, like those elsewhere in the Arboretum, would be valued based upon their full documentation and provenance, not simply a hierarchy of wild over cultivated origin. The attention to cultivated diversity would also solve a problem for us: come midsummer, few things in the garden bloomed. By adding new cultivars of roses (and other species), we could increase the garden's display potential. Around this time, we were dramatically rethinking how we deployed resources for horticulture. Under the leadership of former Deputy Director Richard Schulhof, the Arboretum launched an initial Landscape Management Plan (LMP) in 2007. The LMP prescribed expectations for arboricultural and horticultural care throughout the collections, including the BRC. Simultaneously, we undertook a curatorial review of all plants in the BRC to determine which lineages to preserve. The garden was packed, and the last thing we wanted was to renovate again in ten years. Even so, we still lacked the perspective that only a garden designer could provide. We needed innovative ways to organize plants that would maximize display potential (including for \"BIO\" plants that possess \"botanical interest only\"). We also needed to improve the visitor experience. Without a formal entrance, the garden lacked a sense of arrival. Circulation also demanded attention. While the existing bed configuration (full of tall, dense shrub masses) created intimate garden rooms, it also inhibited visitor exploration due to fear of the unknown or even concerns for safety. In Julie Moir Messervy Design Studio, we found technical expertise to help us tackle these and other problems. Erica and Julie provided both creative genius and sensitivity to the project (and its idiosyncratic client). Although we didn't intend to completely redo the five-acre site, we were seeking a major renovation. Shortly after the project launched in 2008, the Great Recession hit, forcing us to adjust the initial timeline that had called for a single season of renovation using contracted labor. The budget also caused us to rethink aspirations for formal paths and benches, at least initially. Instead, we used our in-house team of horticulturists and interns for the renovation, extending the bulk of the project from one year to three. With JMMDS's new plans in hand, we completely deconstructed some beds, while others were reshaped or constructed anew. This involved handling some 10,000 double cobbles, and a parade of pallets formed along Forest Hills Road. Shifting stones. Double cobbles were installed around a new bed in September 2010, while exposed soil remained visible in the footprint of an old bed (at right). The new plans identified plants to remove, as well as plants that could shift to new locations in the BRC. In some cases, the move could occur instantly, but because of the renovation's phased approach, other plants were transplanted to a temporary location near the Dana Greenhouse and Nursery's south nursery until new beds could be created. This intricate dance kept staff on their toes as they ensured plants were labelled and documented at all times. The greenhouse and nursery staff maintained these, while repropping important accessions from the BRC and beyond. Before we knew it, old beds morphed into turf and new beds appeared. Fresh rose cultivars extended the bloom season in the Rose Roundabout as well as other spaces in the garden. The Prunus Promenade materialized as trees were planted out (including additional cultivars). By the spring of 2011, most of the work was completed, including the installation of Peter Andruchow's beautiful arbor in the Roundabout. This destination for climbing roses was dedicated to Elizabeth Cabot Sluder (daughter of Eleanor Cabot Bradley) during an event with family and staff on June 18. We continue to implement elements of the JMMDS plan for the Bradley Rosaceous Collection. The orchard, comprising a diversity of harder-to-acquire germplasm, has taken some time to fill in, but as of this spring, we are about three-quarters of the way there. In 2013, we hired Peter to construct a second arbor, identical to the one in the Roundabout, which serves as an entry portal between two new beds along Meadow Road. We have also installed granite seating--akin to the bench Julie helped design above the BRC years before--as part of the Arboretum's new Commemorative Bench Program. Gardens, ever changing and dynamic, evolve and grow. The nearly 150-year history of this location, near the Forest Hills Gate, is a perfect example. Michael S. Dosmann is Keeper of the Living Collections and Stephen Schneider is Director of Operations at the Arnold Arboretum. MICHAEL DOSMANN MICHAEL DOSMANN Bradley Rosaceous Collection 11 "},{"has_event_date":0,"type":"arnoldia","title":"Shrubs and the Pollinators Who Love Them","article_sequence":3,"start_page":17,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25642","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15e816b.jpg","volume":75,"issue_number":4,"year":2018,"series":null,"season":null,"authors":"Walker, Holly; Gagliardi, James","article_content":"Shrubs and the Pollinators Who Love Them James Gagliardi and Holly Walker P From butterflies and bees to beetles and birds, many different kinds of pollinators have evolved within their ecosystems, building unique relationships with plants. When gardeners think of designing landscapes for pollinators, they may imagine plantings of floriferous herbaceous beds; however, trees and shrubs are essential components of the habitat required to support a wide variety of pollinators. For successful pollination, a pollinator must find a flower with a structure that matches its body. Consider a butterfly feeding on a daisylike composite flower. The butterfly will gracefully land on the inflorescence and elegantly unfurl its proboscis, which it precisely inserts through the long narrow tube of a central disc WILLIAM (NED) FRIEDMAN ollinators are an essential part of our gardens, the ecosystem, and the United States economy. One in three bites of food you eat depends on pollinators. Honeybee pollination adds more than $15 billion to the value of agricultural crops in the United States each year, with another $9 billion coming from pollination by other species. Pollinator populations have been declining after decades of stress related to loss, degradation, and fragmentation of habitat; reduction in the number and quality of food sources; a lack of sites for breeding, nesting, and roosting; and improper use of pesticides and herbicides. Gardeners can be part of the solution to pollinator loss by creating landscapes that support pollinator health. This congregation of honeybees (and one beetle) was spotted on bigleaf magnolia (Magnolia macrophylla ssp. macrophylla, 961-89*B). DANNY SCHISSLER 18 Arnoldia 75\/4 ? May 2018 Pollen loads the hind legs of this honeybee, which was noticed on the early spring flowers of an Ozark witch-hazel (Hamamelis vernalis, 6099*D). flower to drink nectar hidden inside. During this process, a cleverly positioned anther (male flower part) rubs against the butterfly depositing pollen. After drinking nectar, the butterfly flutters away to the next bloom where the pollen will be brushed against the stigma (female flower part). Now consider beetles, which are sometimes referred to as \"mess and soil\" pollinators because of how they blunder their way through blossoms searching for food. Beetles are important pollinators for flowers like those of magnolia species, often arriving early in the season when temperatures are still too cool for most other pollinators. Since beetles did not originally evolve as pollinators, plants had to adapt to find a way to lure these insects. Most earlyseason beetles are attracted to rotting materi- als, as many beetle grubs are decomposers of decaying wood and plant tissue. The strongly fruited or slightly fetid smells associated with magnolias play on these preferences. Beetles don't possess special pollen-collecting features. Instead, with magnolias, they get covered in pollen while chewing on anthers and tepals (the term for undifferentiated petals and sepals). They then carry the pollen to the next flower. The process may not be as refined as that of a butterfly, but it is just as necessary for certain plants. Beetles are ancient and rank among the earliest evolving pollinators. Therefore, their correlation as the pollinator of ancient plants like magnolias makes evolutionary sense. Beetles and magnolias existed before bees and butterflies, and though bee pollination has been observed on several mag- JAMES GAGLIARDI Pollinators 19 At Smithsonian Gardens, the Urban Bird Habitat is located on the south and west sides of the National Museum of Natural History, not far from the Pollinator Garden. JAMES GAGLIARDI occupies a relatively small footprint in the urban landscape. In a 400-by-40-foot space, we cultivate more than two hundred types of plants, including many woody species. Diversity and seasonality are among the most important factors when choosing plants for pollinators. Plants with high wildlife value and great aesthetics help our public landscape achieve our goals of creating a healthy ecosystem, while both attracting and educating our visitors. Selecting Shrubs for Pollinators Virginia sweetspire (Itea virginica `Henry's Garnet') is an exceptionally adaptable shrub in the Smithsonian's landscape. nolia species at the Arnold Arboretum (see Juan Losada's 2014 article in Arnoldia), the unique connection between these beetles and plants was developed well before other players arrived to the pollination game. At the Smithsonian Pollinator Garden in Washington, DC, we showcase trees, shrubs, and herbaceous plants that support a wide array of pollinators and other wildlife. The garden The evolution and lifecycle of shrubs make them a particularly important part of a wildlife garden. All of the shrubs on this list are tough and adaptable in Mid-Atlantic gardens and often beyond. In addition to producing beautiful pollinator-attracting blooms, many have multi-season horticultural impact. When choosing the best combination of plants for the landscape, be sure that you plan for a garden that serves pollinators throughout the seasons. Include shrubs that flower early in spring, as well as others that will attract heavy pollinator activity in summer or during the KYLE PORT Large fothergilla (Fothergilla major, 968-88*A) attracts pollinators from April to May, much like its smaller-statured relative (F. gardenii). Virginia sweetspire (Itea virginica, Zone 5?9) is another versatile spring-blooming shrub in the Smithsonian's landscape. It is prominently featured at the entrance to the Pollinator Garden in the shade of a black gum tree (Nyssa sylvatica, Zone 3?9). We have also used it in a full-sun planting, where it retains a slope along a parking lot. It thrives in both locations, but the blooms are best in full sun. Its spires of fragrant white blossoms appear in early summer and draw nectar-loving insects like butterflies and bees, including native bumblebees and sweat bees. Best planted in a mass, the plant's rich red to purple fall color will persist into the winter in southern areas as a semi-evergreen plant. If some of the foliage has been damaged by insects, it might be a sign of more pollinator activity, as this plant is a host to the American holly azure butterfly (Celastrina idella). Dwarf fothergilla (Fothergilla gardenii, Zone 4?8) has showy bottlebrush inflorescences that emerge from March to May before their leaves. They attract bees with their white color and pleasant scent, and then offer their pollinators a sweet nectar reward for those that get past their dense tangle of anthers. Ornamentally, it is a great three-season plant with nice foliage that becomes particularly attractive in fall, when it turns brilliant shades of yellow, orange, and red. Consider pairing fothergilla with oakleaf ANSEL OOMMEN, BUGWOOD.ORG DANNY SCHISSLER 20 Arnoldia 75\/4 ? May 2018 Japanese spicebush (Lindera obtusiloba, 376-86*A) produces early spring flowers like its American counterpart (L. benzoin). bustling fall when pollinators are building their winter reserves. Witch-hazel species bookend the pollination season in our gardens. Common witch-hazel (Hamamelis virginiana, USDA Hardiness Zone 3?8) is one of the last plants that will bloom each year. When the days grow short and little else is flowering, the strap-like petals and strong fragrance of witch-hazel flowers draw pollinators like owlet moths, and potentially gnats and late-season bees scavenging for food. At the start of the year, Ozark witch-hazel (Hamamelis vernalis, Zone 4?8) is one of the first blooms to greet pollinators. The caterpillars of spicebush swallowtail (Papilio troilus) feed predominantly on the foliage of northern spicebush (Lindera benzoin). Pollinators 21 JAMES GAGLIARDI In 1995, Smithsonian Gardens opened the Butterfly Habitat Garden, along the east side of the National Museum of Natural History in the heart of Washington, DC. After twenty-one years, this popular landscape was rededicated as the Pollinator Garden. The new theme helps visitors discover the who, what, when, where, why, and how of pollination by interpreting the unique relationship between pollinators and flowers. The garden's title change and our extended educational efforts reflect the growing importance of supporting pollinator health (not just butterflies alone), as championed with a task force formed by President Barack Obama in 2014. Furthermore, the garden is part of the Million Pollinator Garden Challenge, launched by The National Pollinator Garden Network. This effort is a partnership between conservation organizations, gardening groups, volunteer civic associations, and participating federal agencies. It aims to inspire people and organizations to create more pollinator habitats by registering a million public and private gardens and landscapes that support pollinators. As a key advocate for pollinators, the Smithsonian's reinterpretation of the Pollinator Garden on the National Mall educates millions of visitors on the wide diversity of pollinators and the types of plants that support them. JAMES GAGLIARDI The Pollinator Garden James Gagliardi, below, helped with redeveloping the Pollinator Garden to showcase an evolving national awareness of the importance of all pollinators. The garden was officially dedicated in June of 2016. hydrangea (Hydrangea quercifolia, Zone 5?9), which likes similar growing conditions along a woodland edge. Its flowers appear from May to July, after fothergillas have finished, and draw later-emerging wasps and flower flies, along with the aforementioned bees. Not all plants will contribute as much to the aesthetics of your landscape as they will to pollinators. Spicebush (Lindera benzoin, Zone 5?9) is not often the focal point in a garden, but it offers a full package of horticultural benefits. It is tolerant of deer, drought, heavy shade, and clay soil. Green-yellow flowers appear in early spring before leaves emerge, and while the flowers are small, they have garnered enough attention for this native woodland understory shrub to be called the \"forsythia of the wilds.\" The plants are dioecious, requiring small bees and JAMES GAGLIARDI 22 Arnoldia 75\/4 ? May 2018 The drupes of northern spicebush (Lindera benzoin) are popular with bird species, including gray catbird (Dumetella carolinensis), snacking here even before the fruits were completely ripe. various flies to move pollen from the larger, showier flowers on male shrubs to those on the separate females, providing a critical resource for native pollinators when many food sources are not available on the landscape. Once pollinated, the female shrubs produce red drupes that are a good food source for birds and a possible nutmeg substitute for bakers. The plant also features aromatic leaves that turn an attractive yellow in the fall. Spicebush is one of the few host plants used by the spicebush swallowtail (Papilio troilus), a well-recognized visitor of gardens and natural landscapes in the eastern United States. The adult female spicebush swallowtail has evolved to recognize specific compounds on the surface of its host plant before laying eggs, to ensure a suitable food source for maturing larvae. Other related hosts to spicebush that can support the native spicebush swallowtail include sassafras (Sassafras albidum, Zone 5?9) and redbay (Persea borbonia, Zone 7?11). Red chokeberry (Aronia arbutifolia, Zone 4?9) supplies a great deal of value to wildlife in our gardens. Butterflies and native bees, such as mason bees, mining bees, and bumblebees, visit its clusters of white flowers from March to May. Chokeberry foliage turns stunning shades in fall and provides a food source to some hairstreak butterflies and moths, including bluish spring moths (Lomographa semiclarata) and praeclara underwings (Catocala praeclara). This herbivore activity can be observed as typical chewing damage along leaf margins. True to their name, red chokeberry fruits (though actually pomes and not berries) have a dry, astringent taste for birds and humans alike. The fruits persist from summer into the winter, and after a long period of exposure to cold weather, the fruit becomes more palatable. This makes chokeberries an important late-season native food for birds after other food sources are exhausted. The fruit's persistence through late winter also makes it a beautiful ornamental plant in the winter JAMES GAGLIARDI SUSAN HARDY BROWN Pollinators 23 KYLE PORT Attractive throughout the seasons, red chokeberries (Aronia arbutifolia) flower in mid-spring and bear lustrous red fruits well into winter. Flowers of inkberry holly (Ilex glabra `Compacta', 745-69*D) are tucked within dense evergreen foliage. garden. Both red chokeberry and black chokeberry (Aronia melanocarpa, Zone 3?8) are useful for mass plantings or for mixing into a naturalized perennial border. Running serviceberry (Amelanchier stolonifera, Zone 4?8) is another plant that provides showy white blooms for pollinators in May, edible berries in summer, and striking fall foliage. Hollies (Ilex spp.) are a strong draw for pollinators in late spring and early summer. Their small scented flowers often go unnoticed by garden designers, but they effectively draw the attention of bees and flies. The prolific and showy fruits of winterberry holly (Ilex verticillata, Zone 3?9) are another credit to good cross-pollination required for these dioecious plants. At the Pollinator Garden, we recently installed a grouping of dwarf American holly (Ilex opaca `Maryland Dwarf', Zone 5?9), which provides the classic evergreen holly appearance in a more compact space. Other hollies like inkberry (Ilex glabra, Zone 4?9) may not have showy flowers or fruits, but you'll still find them covered in bees and later with birds looking for a snack. The same is true for the common wax myrtle (Morella cerifera, Zone 7?10). Shrubs can also fill a flowering lag in the summer landscape. Buttonbush (Cephalanthus occidentalis, Zone 5?9) 24 Arnoldia 75\/4 ? May 2018 Pollinator Syndromes The combinations of floral characteristics associated with particular types of pollinators are known as pollinator syndromes. Other than bat pollination, which most often occurs in tropical and desert ecosystems, all of these syndromes can be observed in the Smithsonian Pollinator Garden. The Smithsonian team has adapted this information to create seven \"pollinator profiles\" for bees, beetles, butterflies, hummingbirds, flies, moths, and wind (along with special references to bats and water). Using a field-journal theme, each profile in our Pollination Investigation describes the pollinators' favorite flowers based on floral characteristics. The panels teach pollination on a general level and are not designed for our garden alone. On May 20, 2018--the United Nation's first World Bee Day--the panels were unveiled at University of Ljubljana Botanical Garden in the Republic of Slovenia, in recognition of Slovenia's leadership proposing the new celebration. Additionally, our Pollination Investigation panels are available to educators in gardens around the world free of charge through the Smithsonian Gardens website. USDA FOREST SERVICE POLLINATOR SYNDROME TRAITS TRAIT BATS BEES COLOR Dull white, green, or purple Bright white, yellow, blue, or UV NECTAR GUIDES Absent BEETLES BIRDS BUTTERFLIES Dull white or green Scarlet, orange, red, or white Bright, including red and purple Present Absent Absent Present ODOR Strong musty; emitted at night Fresh, mild, pleasant None to strongly fruity or fetid NECTAR Abundant; somewhat hidden Usually present Sometimes present; not hidden POLLEN Ample Limited; often sticky and scented Ample Modest FLOWER SHAPE Regular; bowl-like; closed during day Shallow; have landing platform; tubular Large bowl-like; magnolia Large funnel-like; cups, strong perch support FLIES MOTHS Pale and dull to dark Pale and brown or dull red, purple; purple, pink, flecked with or white translucent patches WIND Dull green, brown, or colorless; petals absent or reduced Absent Absent Absent None Faint but fresh Putrid Strong sweet; emitted at night None Ample; deeply hidden Ample; deeply hidden Usually absent Ample; deeply hidden None Limited Modest in amount Limited Abundant; small, smooth, and not sticky Regular; tubular without a lip Regular; small and stigmas exerted Shallow; Narrow tube funnel-like with spur; or wide complex and landing pad trap-like Pollinators 25 JAMES GAGLIARDI DANNY SCHISSLER is a captivating and attention-grabbing plant for sun to part shade. Planted at the entrance to our Pollinator Garden, buttonbush draws the interest of visitors with its unique round flower heads. The flowers are also a magnet to bees and butterflies in June, just as our hot DC summers begin to peak. Additionally, the flowers of American beautyberry (Callicarpa americana, Zone 6?10) often go unnoticed by gardeners in June and July but draw bees and butterflies for pollination, enabling the glossy purple fruit that gardeners and birds adore. In full sun, bluebeard (Caryopteris ? clandonensis, Zone 6?9) and leadplant (Amorpha canescens, Zone 2?9) are other great summer pollinator plants and can mix nicely into a perennial border due to their smaller habits. Carolina allspice (Calycanthus floridus, Zone 4?9) is also called sweetshrub and strawberry bush because of how the bloom fragrance combines hints of pineapple, strawberry, and banana. Similar to magnolias, Carolina allspice has tepals and evolved long before bees and butterflies entered the landscape. As such, its flowers are predominantly pollinated by sap beetles, though they are attractive to other local pollinators as well. The beetles are drawn by the scent of sweet fermentation, and they work their way into the shade of the overlapping tepals to find food from April to July. The flowers are easy to enter but difficult to depart. Once trapped inside, the beetle picks up pollen. After the flower further matures, the inner parts of the flower fold back to release the beetle. By that point, the stigmas will have already withered, and the beetle will move on to another flower in search of more food, unknowingly ensuring cross-pollination. Summersweet (Clethra alnifolia, Zone 3?9) attracts a diverse group of pollinators, including butterflies, bees, and hummingbirds, which have evolved to take advantage of narrow, tubular flowers. It is one of the few blooms you can find in late-summer shade in our Pollinator Garden. Similarly, bottlebrush buckeye (Aesculus parviflora, Zone 4?8) draws butterflies, bumblebees, and hummingbirds from July to August with its big showy panicles of flowers that occur in part to full shade. Note that the ruby-throated DANNY SCHISSLER 26 Arnoldia 75\/4 ? May 2018 Summer shrubs for pollinators include, top to bottom, buttonbush (Cephalanthus occidentalis, 123-78*A), Carolina allspice (Calycanthus floridus), and summersweet (Clethra alnifolia var. alnifolia `Rosea', 239-47*MASS). Pollinators 27 DANNY SCHISSLER hummingbird (Archilochus colubris) is the only breeding species of hummingbird on the East Coast each summer. It usually arrives in Washington, DC, in April, after migrating north from Mexico and Central America. Summer pollinators also love Chenault coralberry (Symphoricarpos ? chenaultii `Hancock', Zone 4?7). We planted a large grouping of it at the National Museum of Natural History to retain a steep slope, and it may attract the greatest density of pollinators of any shrub in the collection. The plants are thriving and often need to be trimmed, but because they are so popular with honeybees from a hive in the museum's insect zoo, located a few hundred feet away, our gardeners refrain from working with the plants during the summer months, preserving our record of being sting free. Both native and non-native viburnums (Viburnum spp.) work as powerhouses in the landscape, as they attract an exceptionally wide range of pollinators with strong scents that promise either a nectar or pollen reward. Scarab beetles of the genus Cetonia are particularly interesting viburnum pollinators, possessing branched hairs on their bodies that are similar to pollen-collecting hairs found on bees. These hairs ensure a better chance of cross-pollination for self-sterile viburnum species. Beetles, however, are only one of myriad pollinators that are necessary for the successful reproduction of viburnums. As Michael Donoghue reported in Arnoldia in 1980, viburnums with long corolla tubes and sweet scents are more often pollinated by species belonging to the order Lepidoptera, while viburnums with shorter corolla tubes and muskier odors receive frequent visits from flies and small bees. This relationship corresponds to the size of the insect mouthparts. It is important to note that most viburnums produce very little nectar despite the wide range of pollinators associated with the genus. It is thought that the primary reward, at least for bees, is not nectar but pollen. More than Flowers Bottlebrush buckeye (Aesculus parviflora, 558-2003*C) produces feathery panicles from July to August. Flowers are not the only consideration when creating a garden for pollinators. We must consider the needs of pollinators throughout their entire lifecycle. Creating a habitat means maintaining gardens that provide shelter and food. At the Pollinator Garden, we wait to cut back plants and remove dead foliage until spring, if at all. To accommodate the full lifecycle of pollinators, we must cater to caterpillars and other immature insects. In Eric Carle's book The Very Hungry Caterpillar, generations of schoolchildren have learned that we will not have beautiful butterflies without munching caterpillars. Caterpillars can be picky eaters, so we plant a wide variety of host plants in the Pollinator Garden. Pollinators often rely on specific trees, shrubs, perennials, and annuals as JAMES GAGLIARDI food sources. Some plants, like spicebush (Lindera benzoin), the host plant of the spicebush swallowtail butterfly, have pollinator-friendly flowers. But even wind-pollinated species can be important for pollinators. The foliage of smooth alder (Alnus serrulata, Zone 4?9), for instance, provides a significant food resource for beetles, aphids, moth caterpillars, and other insects. When planting a garden for pollinators, we need to be okay with leaves being eaten. It is also best to acquire plants from nurseries that have not treated their plants with systemic insecticides. In the 180 acres maintained by Smithsonian Gardens, we do use insecticides, but only as a last resort. Our preferred methods of control are mechanical, cultural, and biological. The plants in the Pollinator Garden are in good health, in part because maintaining a diverse plant inventory supports a balanced garden ecosystem. During our tenures at Smithsonian Garden, neither of us can recall spraying insecticides in the Pollinator Garden or in the preceding Butterfly Garden. In the extreme case that we ever need to apply an insecticide in the future, we would certainly make sure that the Horticultural care is an important factor when gardening for pollinators. At the Pollinator Garden, horticulturists wait to remove winter foliage, which is product would not affect beneficial necessary for insect habitat. insects and pollinators. In the end, pollination is all about survival James Gagliardi is a supervisory horticulturist with and sex. The insect and the plant both require Smithsonian Gardens in Washington, DC. After President something. The pollinator is often drawn to a Obama released a memorandum to promote pollinators plant with an offer of food. In turn, the plant in 2014, he worked on a task force with the Council on uses the pollinator as a vector to move its Environmental Quality to draft Supporting the Health of Honey Bees and Other Pollinators. He is honored to pollen to the stigma of another flower. Plants be the editor of the Smithsonian's first gardening book, have evolved with particular traits, and polEncyclopedia of Garden Plants for Every Location. linators select blooms for their preference for Holly Walker is the Plant Health Specialist at color, odor, nectar, nectar guides, pollen, and Smithsonian Gardens in Washington, DC. With a diverse flower shape. These traits, combined with background in integrated pest management (IPM), bloom period and location, make for a variable biological control, and native pollinator conservation, matrix of pollinator and plant interactions. she works to educate the public in environmentally Therefore, it is important to grow a large selecresponsible pest management in both urban and tion of plants, including shrubs, to support the rural landscapes. She recently completed her PhD in entomology at the University of Delaware. needs of a great variety of pollinators. JAMES GAGLIARDI 28 Arnoldia 75\/4 ? May 2018 "},{"has_event_date":0,"type":"arnoldia","title":"2017 Weather Summary: A Year of Recovery","article_sequence":4,"start_page":29,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25640","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15ebb6d.jpg","volume":75,"issue_number":4,"year":2018,"series":null,"season":null,"authors":"Pfeiffer, Sue A.","article_content":"2017 Weather Summary: A Year of Recovery S. A. Pfeiffer A DANNY SCHISSLER s an Arboretum horticulturist, my daily routine always starts with the weather forecast. The forecast dictates priorities and deadlines for horticultural tasks. As the forecast shifts, new goals arise and others are eliminated. Pests and diseases come and go with seasonal turns in the weather, and access to our plant collections is determined on a daily basis, especially in the wet and muddy weeks of spring. Past weather conditions, however, also play an important role in directing our horticultural care. By the start of meteorological winter in December 2016, concern over the health of the collections was mounting. Following an exceptionally dry 2015 season, record-breaking heat and drought made 2016 an especially tough Spring colors swept down Oak Path on May 11. year for plants. From June through October, hundreds of feet of hose were dragged through the collection, and every water tank on hand was filled and refilled in an effort to alleviate drought stress. Water cannons pumped six million gallons of water into the landscape, and irrigation systems were on a tight overnight schedule. The efforts of the horticulture crew to carefully monitor and provide supplemental irrigation in the searing heat of 2016--the hottest summer on record for the city of Boston (but the second hottest according to Arboretum data)--cannot be overstated. As we entered 2017, what would it take to end the drought? Temperatures warmed during the winter months, and according to thirty-year averages calculated by the National Oceanic and Atmospheric Administration, precipitation during January and February of 2017 was well above normal for the Boston area. Due to the severity of the ongoing drought, expectations of major S. A. PFEIFFER 30 Arnoldia 75\/4 ? May 2018 Abundant rain brought abundant turf. Arboretum gardener Brendan Keegan is pictured mowing on Peters Hill in early May. DANNY SCHISSLER 2017 Weather 31 Our 218-day growing season enabled vibrant growth throughout the Arboretum. This view shows the North Meadow in mid-August. plant dieback and death were on the forefront of our minds as we awaited spring leaf out. Abundant rains fell during March and April, amounting to more than five inches above normal. Buds swelled, leaves emerged, and a green landscape was a welcome sight. Rains continued to fall. Soils were plenty moist, and the recovery of the collections was exceptional. Not only did the accessioned trees, shrubs, and vines flourish during the growing season but turf and weeds also seemed to grow exponentially, keeping horticulture crews busy. Arboretum staff, always attentive to the impact of weather on plant health, could not have asked for a better year following the preceding drought. Our annual weather summary tracks the four meteorological seasons and reveals, at least in part, moments when weather demanded adjustments in horticultural care. For ease of interpretation and statistical analysis, meteorological seasons are broken into three-month periods based on annual temperature. Winter is defined as the three consecutive months with the lowest average temperatures, corresponding with December, January, and February in the Northern Hemisphere. Spring, summer, and autumn follow accordingly, each comprising the next three-month series. Winter: December 1, 2016 to February 28, 2017 December was seasonable with slightly mild temperatures. Highs dropped below freezing on only four occasions. An outbreak of arctic air on the fifteenth and sixteenth forced temperatures to a low of 3?F, and on the seventeenth, we received five inches of snow. Most melted away as conditions rebounded to 57?F on the eighteenth. The majority of precipitation fell as rain during seven major events. The month of January brought unusual warm conditions. Despite a three-day cold spell that brought single-digit temperatures from the seventh through the 32 Arnoldia 75\/4 ? May 2018 KYLE PORT ninth, January's average temperature was 7.1?F above normal. We hit a recordbreaking 62?F on the twelfth, far above the average high of 36?F for that day. Precipitation was abundant throughout the month with seventeen days of recorded rain or snow. Of this, 3.65 inches fell as rain. Another 9.8 inches arrived as snow, which mostly fell during the three-day cold spell. Major rain events were long, steady, and light, allowing moisture to infiltrate into the soil and recharge groundwater levels. A high-powered coastal storm arrived on the evening of the twentythird, bringing prolonged rain over the next twenty-four hours. Gusts reached 39 mph, scattering limbs throughout the landscape and completely destroying an oak and a willow. Overall, the warm and snowless month allowed access into more remote areas, providing opportunities for horticulture teams to prune deadwood in the beech collection and thin trees in Central Woods. Warm conditions continued into February, as temperatures averaged 6.1?F above normal. The horticulture crew took advantage of snowless days early in the month by pruning and rejuvenating most accessions in the Bradley Rosaceous Collection. We ultimately received above-average precipitation, including 19.2 inches of snow that mostly fell between the seventh and the thirteenth. A nor'easter blizzard, the first since January 2015, delivered 11.5 inches of this total on the ninth. Conditions remained cold and cloudy in the immediate aftermath. With this deep snow cover, horticultural priorities shifted to scouting for signs of invasive insects in the collection. A warm spell sent temperatures soaring between the twenty-third and the twenty-fifth. We reached 74?F on the twentyfourth, the highest February temperature ever recorded in Boston since recordkeeping began in 1872. Daily records were also hit on the neighboring days (69?F on the twenty-third and 72?F on the twenty-fifth). All remaining snow melted The horticulture crew began mulching the Bradley Rosaceous Collection in late January. DANNY SCHISSLER 2017 Weather 33 Chinese witch-hazel (Hamamelis mollis, 215-2000*A) was spotted flowering on February 23. over this period, and the horticulture crew returned to the rose collection, mulching all sixteen beds. Early spring blooms appeared on red and silver maples (Acer rubrum and A. saccharinum) and on hybrid witch-hazel cultivars (Hamamelis ? intermedia `Arnold Promise', `Diane', and `Jelena'). Spring: March 1 to May 31, 2017 March was colder than normal with an average temperature below both January and February. Early in the month, lows dipped into single digits on three occasions. Precipitation was abundant and consistent throughout the month, although we experienced some of the driest air of the year between the third and sixth, when relative humidity levels remained in the teens and low twenties. On the fourteenth, a late-season blizzard brought high winds, gusting at 47 mph, our highest of the year. Heavy snow amounted to 6.5 inches and turned to heavy rain that fell at a rate of 0.50 inches per hour. Temperatures plummeted to the teens overnight, turning roads and sidewalks into a veritable skating rink by the morning of the fifteenth. The storm brought down limbs in the conifers and toppled an apricot (Prunus armeniaca). Temperatures remained cool until vernal equinox on the twentieth brought sunny conditions and a high of 51?F. We ended the month with plenty of rain. Buds that had begun to swell due to warm February temperatures suspended their development, waiting to open, while cooler temperatures extended the bloom time of many witch-hazels. The lack of snow cover allowed the horticulture crew to pursue diverse projects: mulching the beech collection, installing new paths, removing invasive plants in natural areas, and cleaning winter storm damage (including a giant willow that was pulled from the meadow). DANNY SCHISSLER 34 Arnoldia 75\/4 ? May 2018 KYLE PORT Always among the first to brave the spring, Zen magnolia (Magnolia zenii, 430-91*A) was zapped by cold weather on March 5. Abundant rainfall continued into April, further reducing the water deficit from 2016. Temperatures were above average for the month. A storm that lasted from March 31 to April 1 delivered 2.82 inches of precipitation, most falling as rain. We began the third with our last spring frost, which melted as temperatures warmed to 60?F later that day, marking the beginning of the growing season. Excessive rain and melting snow saturated soils, especially in lowlying areas, leaving those areas inaccessible. Ponds filled, brooks flowed, and the forsythia began to bloom. Nursery digging for spring plantings began on the tenth. Warm conditions persisted as we hit 86?F on the eleventh and sixteenth, causing katsuras (Cercidiphyllum japonicum), magnolias (Magnolia), and cherries (Prunus) to burst into flower. Despite seasonal dreary conditions, the landscape looked alive as turf greened up and trees leafed out. Horticulture crews were busy with mowing Low-lying portions of the Arboretum, including the lindens, flooded as the growing season began in early April. 2017 Weather 35 Pests and Pathogens Mind the Weather Too MENG LI MENG LI Not all collections escaped the 2016 drought unscathed. Oaks on Peters Hill, defoliated for many years by winter moth (Operophtera brumata), were among those that continued to languish, as were hemlock specimens that had been struggling from infestations of hemlock woolly adelgid (Adelges tsugae) and elongated hemlock scale (Fiorinia externa). Effects of drought stress on older beeches became apparent mid-summer as leaves browned and black-timber bark beetles (Xylosandrus germanus) invaded. Royal azaleas (Rhododendron schlippenbachii) in the Explorers Garden experienced major stem dieback. The arborist crew devoted a month to removing deadwood from older oaks and other large trees, while horticulturists did the same in the understory. Both powdery mildew and anthracnose, a general term for many fungal leaf diseases, erupted throughout the grounds. From the Arborway to Peters Hill, from May through October, anthracnose was ubiquitous due to frequent precipitation, overcast skies, and warm temperatures. Sycamores (Platanus), maples (Acer), and dogwoods (Cornus), which are typically affected Crabapples on Peters Hill presented an extraordinary spring display, including Magdeburg crabapple (Malus ? magdeburgensis, 544-66*A) and Chinese pearleaf crabapple (Malus asiatica, 341-86*B). by anthracnose, were especially hard hit. Other deciduous trees, including ashes (Fraxinus), beeches (Fagus), hophornbeams (Ostrya), hornbeams (Carpinus), oaks (Quercus), walnuts (Juglans) and stewartias (Stewartia), were certainly not immune. Severe infestation on sycamores and crabapples caused premature defoliation. Fortunately, however, winter moth populations were greatly reduced. Warm temperatures in January and February led to early bud break for red and silver maples (Acer rubrum and A. saccharinum), simultaneously causing winter moths to hatch early. The newly emerged caterpillars could wiggle between bud scales on those maple species and begin feeding, but subsequent cold temperatures delayed bud swelling for later-flowering host plants, notably apples and crabapples (Malus). With no access to the inner buds of those species, many caterpillars didn't survive through the spring. In recent years, Arboretum staff have also released tachinid flies (Cyzenis albicans) that prey on winter moths. The combination of these biotic and abiotic factors significantly lowered the population of winter moths, allowing the crabapple collection on Peters Hill to put on its best flowering show in years. ANDREW GAPINSKI operations throughout the grounds. We ended the month with temperatures hovering just above 80?F, pushing lilacs (Syringa) into bloom. May was slightly warmer than average, but we began the month with below-seasonal temperatures and lows in the thirties. A number of fastmoving downpours accounted for over an inch of rain during the first week. According to the United States Drought Monitor, soil moisture returned to normal conditions Arboretum horticulturists Mark Walkama (left) and Scott Grimshaw prepare a ball-and-burlap specimen in the East Nursery. This on May 9, officially ending (right) year, 378 plants graduated into the collection. the drought that had begun on June 7, 2016. Overcast conditions prevailed for the week leading up to Lilac Sunday on the fourteenth. These cooler temperatures extended the blooms of many plants, especially lilacs, but with the threat of soaking rain on the fourteenth, Lilac Sunday activities were held on Saturday the thirteenth. True to forecast, a nor'easter arrived that evening, and by the afternoon of the fourteenth, we had received over 1.5 inches of rain. Temperatures soared over the following days, hitting highs in the nineties from the seventeenth to the nineteenth and peaking ANDREW GAPINSKI 36 Arnoldia 75\/4 ? May 2018 Arboretum arborists worked along Meadow Road on May 8, the day before the United States Drought Monitor officially announced an end to the drought that began on June 7, 2016. DANNY SCHISSLER 2017 Weather 37 Japanese pagoda tree (Styphnolobium japonicum, 216-35*A) showcased the lush greenness of this recovery year, pictured on August 2. at 96?F. This heat wave was one of the earliest for the area, as the most recent with an earlier date occurred May 2?4, 2001. A fast-moving cold front brought a heavy downpour on the evening of the nineteenth: within fifteen minutes, 0.30 inches of rain had fallen and temperatures had plummeted by 10?F. We ended the month with typical spring weather fluctuations and steady precipitation. Soils remained moist, plants were lush and floriferous, and turf continued its rapid growth. Horticulture crews were busy mowing and finishing spring cleanup. Summer: June 1 to August 31, 2017 A lingering cold front in early June brought temperatures in the fifties. Four days of rain accounted for 2.0 inches of precipitation, almost half of the monthly total. Moist soil conditions made it difficult to access low-lying areas for mowing operations. We experienced our second heat wave of the year between the eleventh and thirteenth, once again hitting 96?F. High soil moisture and hot temperatures led to an explosion of turf and weed growth throughout the grounds. The plant collections flourished, easing concerns over last year's drought as plants continued to recover. A fast-moving system brought an additional 1.78 inches of rain on the evening of the sixteenth. Conditions stabilized as temperatures remained in the high seventies to mid-eighties for the remainder of the month. Drier conditions were prevalent, and despite seven short rain events, we accumulated only one additional inch of rain. Nonetheless, compared to June 2016, we received almost four times more rain in 2017. July temperatures were seasonable with slightly below-average precipitation. We began the month with comfortable conditions, highs in the eighties and lows in the high fifties. Two fast-moving thunderstorms on the seventh and eighth brought downpours and a total of 0.94 inches of rain. Thunderstorms returned 38 Arnoldia 75\/4 ? May 2018 What Factored into the Drought Recovery? DANNY SCHISSLER The Arnold Arboretum's primary weather station is located at the Weld Hill Research Building, where it takes measurements for temperature and precipitation every fifteen minutes. Snow measurements are taken once a day at the Dana Greenhouse. In 2017, three environmental factors played key roles in drought recovery for the plant collections. First, precipitation was plentiful in 2017, with a monthly average of 3.98 inches, compared to 2.84 inches in 2016 and 3.09 inches in 2015. Excess precipitation during the first six months of 2017 caused the United States Drought Monitor to announce that Boston had reached normal conditions by early May. During that time, light- to moderate-intensity rainfall occurred frequently. These conditions sustained soil moisture, allowing roots to consistently uptake water for plant growth and metabolic processes. Second, temperatures were mild from January through February and remained seasonable throughout subsequent months, except for a warm spell in October. While New England heat waves typically occur between July and September, when soil moisture levels are predictably low, two of the three heat waves that occurred in 2017 arrived early: one in May, another in June. During these heat waves, plants were not set back to the extent normally expected. Precipitation during both months was greater than expected, supplying relief from the effects of high temperatures. Likewise, despite normal high temperatures during summer months, frequent rain minimized plant stress. Third, we experienced an exceptional 218-day growing season, beginning April 3 and ending November 8. (The growing season is typically defined as the number of days between the last spring frost and the first in the fall.) The season was more than three weeks longer than the average growing season since 2009. Even more astonishing, it was thirty-five days longer than 2016. Moreover, because severe and extreme drought conditions persisted from July 26, 2016, through the first fall frost, the actual period of active plant growth that year was far less than the 183 days recorded between frosts. The Linda J. Davison Rhododendron Path was festooned with lavish pinks on June 5. 2017 Weather 39 Arnold Arboretum Weather Station Data ? 2017 Avg.Avg.Avg.Max. Min. Precipi- SnowMax. Min. Temp.Temp.Temp.tation fall (?F)(?F)(?F)(?F)(?F) (inches) (inches) JAN 40.927.134.061.8 4.7 4.27 9.8 FEB 45.427.136.274.2 9.1 3.8119.2 MAR 41.325.133.263.0 7.8 5.5211.1 APR 61.242.952.186.430.1 5.66 1.2 MAY 63.847.855.895.734.0 4.17 JUN 78.858.168.595.845.7 4.56 JUL 81.862.172.093.052.2 2.82 AUG 80.959.873.390.849.5 2.43 SEP 76.157.666.987.943.0 3.61 OCT 69.550.359.979.437.5 5.96 NOV 51.333.242.275.520.5 1.98 DEC 35.322.028.759.3 0.2 2.9810.4 Average Maximum Temperature . . . . . . . . . . . . 60.5?F Average Minimum Temperature . . . . . . . . . . . . 42.8?F Average Temperature . . . . . . . . . . . . . . . . . . . . . . 51.7?F Total Precipitation . . . . . . . . . . . . . . . . . . . . . . . . 47.77 inches Total Snowfall in 2017 . . . . . . . . . . . . . . . . . . . . 51.7 inches Snowfall During Winter 2016?2017 . . . . . . . . . . 47.5 inches Warmest Temperature . . . . . . . . . . . . . . . . . . . . . 95.8?F on June 12 Coldest Temperature . . . . . . . . . . . . . . . . . . . . . . 0.2?F on December 29 Strongest Wind Gust . . . . . . . . . . . . . . . . . . . . . . 46.9 mph on March 14 Last Frost Date . . . . . . . . . . . . . . . . . . . . . . . . . . . 30.1?F on April 3 First Frost Date . . . . . . . . . . . . . . . . . . . . . . . . . . 29.5?F on November 8 Growing Season . . . . . . . . . . . . . . . . . . . . . . . . . . 218 days Growing Degree Days . . . . . . . . . . . . . . . . . . . . . 3104 days Days at 90?F or Above . . . . . . . . . . . . . . . . . . . . . 11 days Days Below Freezing . . . . . . . . . . . . . . . . . . . . . . 23 days 40 Arnoldia 75\/4 ? May 2018 from the eleventh through the thirteenth, followed by a cold front that brought temperatures in the sixties for a couple days. Temperatures returned to expected highs, and we experienced our third heat wave of the year from the nineteenth to the twenty-first, with temperatures topping out at 93?F. Conditions continued to fluctuate as a slow-moving storm materialized, bringing prolonged soaking rains and temperatures in the sixties on the twenty-fourth and twenty-fifth. Because this addition of 1.47 inches of rain kept soils moist, we had yet to irrigate the collections (except for new plantings). We ended the month with partly cloudy conditions, high humidity, and temperatures in the seventies and eighties. Seasonable temperatures continued in August. High humidity and warm temperatures over the first couple of days culminated in violent thunderstorms that brought heavy downpours, hail, and high winds on the third and the fourth. Almost two inches of rain fell, scattering the landscape with fallen leaves and causing erosion (even in mulched beds) and flooding. A cold front brought conditions in the seventies from the fifth through the eighth, but otherwise, typical New England heat and humidity returned for the remainder of the month. We received only five additional rain events over the remaining four weeks, totaling a mere 0.38 inches. Soils dried, leading to dusty conditions in open grassy areas. Newly established turf began to go dormant, and supplemental irrigation began mid-month. Clear skies were ideal for viewing the partial eclipse, which peaked at 2:48 p.m. on the twenty-first. We reached 91?F on the twenty-second, the single incident of temperatures in the nineties for the month. Although very humid, temperatures were seasonable, remaining in the eighties mid-month and dropping to the seventies during the last week. Serviceberries (Amelanchier), which typically begin to drop their leaves early in the season, remained green and lush throughout the month. Autumn: September 1 to November 30, 2017 September brought average rain that fell predominantly as quick downpours. The month began with temperatures in the sixties and seventies as remnants of Hurricane Harvey delivered a quarter inch of rain on the third. An additional inch arrived as heavy overnight downpours on the sixth, including 0.30 inches that fell over a fifteen-minute period, the first significant rainfall since August 3. Soils were moist and temperatures favorable when autumn planting began on the eleventh. Hot and humid conditions returned mid-month leading to pop-up thunderstorms on both the fourteenth and the fifteenth. Conditions did not improve as Hurricane Jose approached on the seventeenth, bringing dense morning fog that was followed by wind and sporadic rain. Dreary weather persisted with minimal precipitation until the twenty-third, after which the sun and heat returned. Temperatures rose to the mid-eighties between the twenty-fourth and twenty-seventh, 15 to 20?F above average for this time of year. Summer conditions, however, would not last. Temperatures dropped to a more seasonable 56?F when a storm delivered over half an inch of rain on the thirtieth. With the abundance of precipitation in the latter half of the month, irrigation was limited to recent plantings. October was the third warmest on record. Highs shot above 70?F on seventeen occasions, exceeding average monthly temperatures by over 7?F. We began the month with warm and dry conditions, reaching a high of 79?F on the fifth. Two small showers arrived on the eighth and ninth. Warm temperatures continued and precipitation remained minimal. By the seventeenth, we entered moderate MICHAEL DOSMANN 2017 Weather 41 This green ash (Fraxinus pennsylvanica var. subintegerrima `Marshall's Seedless', 626-54*A) numbered among ten trees destroyed during an overnight nor'easter on October 29. drought, mirroring conditions last experienced in late March. Conditions were ideal for the horticulture crew, which spent these weeks seeding renovated areas, clearing the natural area behind the hickory collection, mulching new paths, and planting meadow natives. The dry pattern broke when a slow-moving system dropped over two inches of rain between the twenty-fourth and the twenty-sixth. An overnight nor'easter arrived on the twenty-ninth, bringing heavy rain and strong winds with 37 mph gusts. By the morning, 3.41 inches of rain had fallen and the collections experienced moderate damage; ten trees were lost, and many branches dropped throughout the grounds. Low-lying areas were flooded from the 5.5 inches of precipitation that had fallen over the previous six days, and the drought ended as quickly as it had arrived. We ended the month with highs in the sixties, having yet to receive our first frost. November temperatures were seasonable, despite large fluctuations, and rainfall was below average. We started the month with a continuation of the warm temperatures experienced in September and October, hitting 76?F on the third. These warm temperatures forced a number of spring-blooming shrubs into flower, including Smirnow rhododendron (Rhododendron smirnowii). This would mark the end of an unusually warm autumn; temperatures dropped into the sixties from the fifth through the seventh, before settling in the forties and fifties for the remainder of the month. We finally received a frost when temperatures dipped to 29.5?F on the eighth. This ended our growing season at 218 days. Low temperatures continued to drop as we sunk into the twenties for six straight days, reaching 21?F on the eleventh. These freezing conditions, well lower than expected for this time of year, caused leaves on many trees to freeze and die before abscission cells SUZANNE MROZAK 42 Arnoldia 75\/4 ? May 2018 This specimen of Japanese maple (Acer palmatum var. matsumurae, 148-57*B) was among the maples that carried an abnormal cover of marcescent leaves through the winter. What Happened to Autumn Foliage? Autumn color was poor at the Arboretum due to a series of meteorological factors. Ample moisture and favorable temperatures during the growing season precluded a typical summer drought, encouraging trees to bear lush foliage at the beginning of meteorological autumn. Conditions remained hot and moist through late September and October, and plants showed no signs of slowing down. Because nighttime temperatures remained relatively high, most plants failed to trigger leaf senescence (the final stage of leaf development, which leads to fall color and eventual drop). When arctic conditions descended in November, nighttime temperatures plummeted into the twenties for six straight days. Plants were unprepared for this sudden freeze, which slowed foliage change for some specimens and ended it for others, causing leaves to freeze before developing the abscission layer needed to separate and drop. This phenomenon, called marcescence, left many trees--notably the Asian maples--holding onto their brown, crispy leaves. This slowed leaf drop and prolonged autumn leaf cleanup. DANNY SCHISSLER 2017 Weather 43 Fall color began late and ended with little finale. This view shows leaves in the oak collection. could fully develop. Temperatures remained unseasonably cool with consistent rainfall. The largest storm delivered 0.88 inches of rain on the twenty-second. Sustained cleanup efforts continued throughout the month, as the crew progressed into leaf cleanup after completing storm-damage removal early in the month. Early Winter: December 2017 Cold temperatures extended into December. Highs dropped below freezing on the twenty-sixth and then sunk into the teens from the twenty-eighth through the end of the month--the four coldest days of 2017. Moving Forward The Arboretum experienced an optimal growing season in 2017, yet we cannot close the chapter on the preceding drought before considering the long-term effects of such a prolonged water shortage. Symptoms of persistent plant stress are more often observed years down the road, ultimately causing slow decline and possible death. As plants recover from drought, their ability to defend against disease and insect attacks remains compromised. Bark beetle invasions can be linked to drought stress, as can the onset of Diplodia tip blight and Cytospora and Nectria cankers, but connecting future disease and pest outbreaks to past drought events often proves difficult. Internal plant damage is hidden, and the cumulative effects of long-term drought stress may impact tree health for many years. As we move into 2018 and beyond, vigilance and regular observation will be critical to the overall preservation of the collections. Sue A. Pfeiffer is an Arboretum Horticulturist at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"The Pear to End All Wars: Pyrus ussuriensis","article_sequence":5,"start_page":44,"end_page":45,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25643","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15e856f.jpg","volume":75,"issue_number":4,"year":2018,"series":null,"season":null,"authors":"McDermitt, Matthew","article_content":"The Pear to End All Wars: Pyrus ussuriensis Matthew McDermitt O ne of my favorite things about the annual plant inventory at the Arnold Arboretum--the process of field checking each plant--is that you discover interesting specimens that many overlook. On a windy Halloween afternoon, I encountered a Ussurian pear (Pyrus ussuriensis, accession 11302*C) growing in dense woods along South Street. The tree was struggling for light and space under a canopy of old oaks and would not impress the average observer, but like all plants at the Arboretum, it has a story that is documented in our plant database. When the First World War began in 1914, Ernest Henry Wilson was busy collecting plants in Japan. Charles Sprague Sargent, the Arboretum's director, was travelling in England, and when he realized the gravity of Europe's political conflict, he asked Wilson to cut his trip short. Upon returning to Boston in 1915, Wilson was shocked to learn about the sinking of the RMS Lusitania and the rapid expansion of Central Powers across Europe. At the Arboretum, tensions began to rise between Wilson (an Englishman) and his friend and colleague Alfred Rehder (a German taxonomist). Wilson and his wife, Ellen, began only speaking to the Rehder family in a professional context. Relief came when Sargent sent Wilson on his sixth plant-collecting expedition in January 1917. Just before Wilson's departure, Sargent wrote to a correspondent of \"bad times,\" but he noted that \"there is no use thinking about them when there are trees to think of.\" Wilson spent the next two years collecting plants throughout territory that was then occupied by Japan. In 1918, he collected seed from a Ussurian pear in the Gyeonggi Province of South Korea (then known as Keiki-do, Japan), giving rise to accession 11302. The specific epithet for Pyrus ussuriensis refers to the Ussuri River, which flows from eastern Russia into northeastern China, forming part of the border between the two countries. The species inhabits this region--the coldest and most northern range of any pear species (it is hardy to USDA Zone 3)--along with portions of Korea and Japan. While not of global conservation concern, the species is endangered in Japan. The Arboretum also holds P. ussuriensis var. hondoensis--a Japanese variety, currently listed as vulnerable in Japan--along with the regional cultigens `Pin-li', `Shinchu', and `Shu-li'. The flowers of the Ussurian pear are among the most attractive of the genus: the immature buds are tinged deep red, and the color initially remains as the five petals unfurl. Once fully open, the flowers measure 3 to 3.5 centimeters (about 1.25 inches) in diameter and have beautiful dark red anthers that pop against the white petals. The flowers are similar to those of the Callery pear (Pyrus calleryana), but are notably larger and open earlier in the season. The Callery pear became a wildly popular street tree in North America, but it fell from grace when it escaped into natural areas, not to mention limbs that would drop with the slightest wind. In contrast, P. ussuriensis hasn't demonstrated invasive tendencies or structural flaws. Besides the beauty of the flowers, Sargent was impressed by the size of the species. In the Bulletin of Popular Information, Sargent noted that the Ussurian pear is the tallest and largest of all pear species, citing a specimen that Wilson photographed in Korea, which was 18 meters (60 feet) tall with a trunk diameter of 1.4 meters (4.5 feet). He was also excited that Pyrus ussuriensis appeared resistant to fire blight, a plant disease that plagues many members of the rose family, but unfortunately, minor fire blight has been observed on Arboretum accessions within the past decade. Wilson returned to the Arboretum in early 1919, several months after the First World War had officially ended. Although the original pear that grew from his South Korean collection is hidden in a natural area, an exceptional example of Pyrus ussuriensis var. hondoensis (accession 11728*A) grows on the northern side of Poplar Gate Road. This tree was grown from seed Wilson collected in Nagano, Japan, on the same expedition. Matthew McDermitt is a former curatorial assistant at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23461","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14eb36e.jpg","title":"2018-75-4","volume":75,"issue_number":4,"year":2018,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"2017: A Banner Year for the Campaign for the Living Collections","article_sequence":1,"start_page":1,"end_page":3,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25630","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15ea36d.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Dowell, Robert; Dosmann, Michael S.","article_content":"2017: A Banner Year for the Campaign for the Living Collections Robert Dowell and Michael S. Dosmann S ince the Arnold Arboretum began its Campaign for the Living Collections in 2015, plant-collecting expeditions have launched with a fervor and sweeping extent rarely seen before. So far, 14 separate trips have taken place across Europe, Asia, and North America. These forays yielded incredible plant riches, from new collections of the daintyleaved regal lily (Lilium regale)--first introduced from China to the Arboretum by Ernest Henry Wilson--to the mast harvested from towering American beeches (Fagus grandifolia) in mature forests of eastern North America. With the 2017 season concluded, the Arboretum has progressed well toward its 10-year goal of securing germplasm from almost 400 target taxa or desiderata enumerated in the Campaign's opening communiqu? (Friedman et al., 2016). So far, the Arboretum has acquired 147 taxa from the list, resulting in over 200 accessions now in propagation and production at the Dana Greenhouses and Nursery--with a few already growing in the permanent collections. Deciding how, when, and where to collect a diverse array of target species requires organization, and several precepts guide our efforts. One of these is the \"greatest bang for the buck\" principle: geographic areas with the highest concentrations of plants on our wish list become hot spots for concentrated and repeated expeditionary activity. Our teams focus on these areas first, and as the Campaign progresses and we check targets off the list, high priority areas shift to those of lower priority. Because eastern North America and eastern Asia possess many of our desiderata, they will always remain important collecting spots, as they were in 2017. A second principal in the Campaign gives precedence to species with confined native ranges, or select portions of their native range. For example, the nutmeg hickory (Carya myristiciformis) grows in isolated populations from Texas to South Carolina, and its importance was a reason why a team of explorers went to the coastal southeast. The Campaign also In 2017, Arboretum explorers mounted four major expeditions: three occurred in North America: Wisconsin (WIE), ArkansasMissouri (ARMOE) and the Coastal Southeast (COSE); a fourth was a North America-China Plant Exploration Consortium trip to Sichuan (NACPEC2017). Individual markers are where the explorers made specific collections. MICHAEL S. DOSMANN Campaign 2017 LIVING COLLECTIONS 3 await further trial and selection, such as the paperbark filbert (Corylus fargesii), an up-andcoming horticultural commodity collected on the 2015 NACPEC trip to China. Other collections represent novelty, such as the Georgian oak (Quercus iberica), a species never before grown in the living collections and acquired in 2016 in the Republic of Georgia. Just as we cannot predict all the species we will successfully acquire, we cannot anticipate all of the challenges and hazards encountered along the way. Treacherous road conditions, seasonal drought, and terrestrial leeches are just some of the things that can affect an expedition. Despite these and other encounters in the field, however, the 2017 expeditions bore amazing fruit: over 100 taxa and almost 150 separate collections! This leads to one more reflection about the Campaign for the Living Collections: it is as much about the growth of people as it is about the growth of our collection. This year, eight Arboretum staff members went into the wilds. Whether it was their first or their tenth ? Manager of Horticulture Andrew Gapinski admires a handsome Hydrangea bretschneideri in northern Sichuan during the 2017 NACPEC trip. Although the Arboretum already cultivates three different wild-sourced lineages of this species, the team opportunistically collected a fourth for further evaluation of its ornamental potential. seeks to increase the genetic diversity within certain genera grown at the Arboretum. This means growing as many species as possible within each genus, as well as growing multiple, well-documented representatives of each of those species. This is particularly true for the Arboretum's six Nationally Accredited Plant CollectionsTM: maple (Acer), hickory (Carya), beech (Fagus), Stewartia, lilac (Syringa), and hemlock (Tsuga). For species in these and other priority genera, we seek to grow accessions representing the center and distinct reaches of the native ranges. Beyond acquiring targeted species, the Campaign leverages chance by making supplemental or opportunistic collections in the field. To date, over 350 of these taxa have been collected for the Campaign, many on expedition. Some have promising ornamental merit and + ? ? + The native range of red maple (Acer rubrum) spans much of eastern North America. However until 2017, wild-sourced accessions in the Arboretum's collections were from only three northeastern provenances (?). The 2017 WIE and ARMOE expeditions sought out, and secured, germplasm from two new regions (+). Map modified from: Little, E. L. Jr. (1999). Atlas of United States Trees. U.S. Geological Survey. MICHAEL S. DOSMANN 4 Arnoldia 75\/3 ? February 2018 The 2017 collecting team in Sichuan passed abandoned homesteads and towering Meliosma in their search for rare species. expedition, each collector discovered and grew, and brought those experiences back to the Arboretum. The Campaign articles profiled in this issue of Arnoldia highlight some of those experiences, from the humbling appreciation and celebration of history to the excitement of encountering a familiar species for the first time in the wild. We appreciate how \"reading a habitat\" leads to species acquisition, and how there is no substitute for seeing a plant in the wild in order to figure out how to cultivate it. Lastly, there is the value of persistence: sometimes, to secure a species you must collect it far and wide, and at other times, you must resort to an unusual propagation method. These represent just a few of the stories from 2017, and we look forward to those that the next seven years will bring. Literature cited Friedman, W.E., M.S. Dosmann, T.M. Boland, D.E. Boufford, M.J. Donoghue, A. Gapinski, L. Hufford, P.W. Meyer, and D.H. Pfister. 2016. Developing an exemplary collection: A vision for the next century at the Arnold Arboretum of Harvard University. Arnoldia 73(3):2?18. Robert Dowell was a Living Collections Fellow (2016?2017) and Michael S. Dosmann is the Keeper of the Living Collections at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Keeping the Legacy: Retracing Century-old Footsteps","article_sequence":2,"start_page":5,"end_page":8,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25636","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15eb326.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"LIVING COLLECTIONS 5 PHOTO BY THE AUTHOR Keeping the Legacy: Retracing Century-old Footsteps Michael S. Dosmann A ugust steamed hot as plant explorer Ernest Henry Wilson traversed the wilds of northwestern Sichuan in 1910, leading his fourth expedition to China. At this point in his career, many of the species Wilson saw were becoming familiar to his eyes, even mundane, and he was eager for something different. Although the expedition targeted conifers he observed on previous excursions, namely firs (Abies) and spruces (Picea), he was eager to see new habitats and with them, new species. On earlier trips, Wilson had visited Songpan--his destination--from Chengdu, Sichuan's capital to the south, using one of two routes. The most direct option tracked the Min River north, though it was rather depauperate in botanical diversity. Wilson had also used a counterclockwise route that bore him north-northeast of Chengdu through modern-day Mianyang and beyond to Pingwu (Longan Fu in his day), and then west-northwest to Songpan. This time, he forged a way through an expanse few westerners save a couple of missionaries had braved before. The path cut between the previous routes and would provide him the opportunities he sought. On the outset of this trip's leg, he did indeed encounter several things for the first time: he was robbed, from one of his own porters no less, and a local official dismissed his request for an escort through the unchartered territory. This same region of Pingwu became the destination for the 2017 NACPEC (North America?China Plant Exploration Consortium) expedition, which Andrew Gapinski (Arnold Arboretum), Jon Shaw (Harvard Magazine), Kang Wang and Jian Quan (both from Beijing Botanical Garden), Huaicheng Li (Chengdu Institute of Biology), and I undertook from 15 September to 1 October, 2017. I had tracked Wilson's footsteps several times in the past, quite literally during the 2014 filming of Chinese Wilson (a documentary produced by Earthquakes followed by heavy rains caused many of the roads in Pingwu County to subside into the rivers. The collecting team, and the wandering yak-cow hybrids seen here, choose their footsteps carefully. Central China TV), seeing towns, roadways, mountain views, and of course plants that he had also visited. However, this trip was more poignant, for I now bore the title of Keeper of the Living Collections; the title of Keeper used only once before at the Arboretum, given to Wilson in 1927. During our trip, we endured roadways capsized during the season's earthquakes and rains, and we got up-close-and-personal with terrestrial leeches that slink ubiquitously in these rich, mesic forests. On the morning of September 23, our team set out to explore one of the many mountain valleys near Si'er, an area Wilson referred to as Tu-ti-liang shan or mountain. Wilson had been awestruck by the herbaceous DIGITAL COLLECTION OF THE HARVARD UNIVERSITY HERBARIA 6 Arnoldia 75\/3 ? February 2018 torrent below, we were impressed with the expanses of large trees to either side. We spotted dove tree (Davidia involucrata), several species of birch (Betula spp.) and gargantuan specimens of the multi-stemmed Chinese beech, Fagus engleriana. Giant panda, we learned from our guide, often overwintered between the stout boles of these very beech trees. Wilson's 1907 collection of the beech from Hubei is the only representation at the Arboretum, and likely anywhere else. Thus, on our hike back down the mountain, we secured fruits of this and other species. Wilson may not have recognized the heavily wooded landscape that we saw. He described this sparsely populated hamlet of Hsao-kou (now Xiaogou and depopulated) as having \"... open, park-like slopes, quite unlike anything I have encountered elsewhere in China. Now largely denuded of trees these glades are covered with grass, and horses, goats, and pigs are raised here in some quantity.\" The lumberman's axe and the herder's livestock had eliminated many of the conifers Wilson had hoped to find in the area. However, A fruiting, type specimen (Wilson 4301) from the Herbarium of the Arnold Arboretum (A) of Cercidiphyllum japonicum var. sinense. In his hard-toon August 17, 1910, he enthusiastidiscern handwriting, Wilson notes \"one tree measured 55ft. in girth 5 ft above cally noted an abundance of Cerciground!\" Although the variety is no longer accepted, plants with the pubesdiphyllum japonicum (katsura tree) cent leaves and follicles can be recognized as C. japonicum f. miquelianum. growing throughout the landscape. plants there, in particular the fingerleaf rodgAlthough he had seen the species on his previersia (Rodgersia aesculifolia), which grew \"in ous travels, he had never seen so many growing [the] millions.\" \"It was in the fruiting stage,\" he together. The intrepid explorer got busy with noted in China, Mother of Gardens, \"but when his vasculum, collecting herbarium vouchers. in flower the acres of snow-white panicles must This collection (Number 4301) would reprehave presented a bewitching sight. Nowhere sent the type of what he and Alfred Rehder later else have I seen this plant so abundant or luxudescribed as a separate botanical variety with pubescent leaves and follicles, Cercidiphyllum riant.\" In an adjacent valley the previous day, japonicum var. sinense. we had seen the same species, each of its coarse Wilson reported trees up to 36.6 meters (120 leaflets reaching nearly a half-meter (20 inches) feet) tall, and 2.1 to 6.1 meters (7 to 20 feet) in in length. As enthralled with it as Wilson had girth. Even amidst these giants, one specimen been, we joyously made a collection. stood out for its enormity--not in height but in As we ascended the valley on foot, marching girth: it was multi-stemmed and hollowed in along a roadway that was tumbling into the JONATHAN SHAW\/COURTESY HARVARD MAGAZINE the core, yet 16.8 meters (55 feet) around, which would be a diameter of 5.3 meters (17.5 feet)! Words are not necessary to conjure what Wilson saw, for he memorialized the tree in a photograph. This exact tree was our destination for the day's hike, and for me it glimmered with personal significance, because Cercidiphyllum japonicum had drawn me to China for the first time in 1999 and I considered this a bit of a reconnection. This was now my eighth trip to China, and perhaps a bit like my predecessor, I sought some reprieve from the mundane. I imagine that due to the openness of the landscape 100 years ago, Wilson spied the specimen easily from the path. Yet, even after our team left the road and ascended the steep and muddy streambanks, dense woods left us completely blind to our target. We had to bushwhack through the brush, and upon arriving at last, we could do nothing but marvel. Like the forest around it, the tree had recovered over the last century. The specimen had been but 8 meters (25 feet) tall in 1910; it now stood over 20 meters (65 feet) in height, and the diameter of the largest stem was just under 2 meters (6.5 feet). The three massive, original basal stems remained, though showed considerable wear. The tree at Wilson's time was probably coppiced repeatedly for firewood, and it had since rebounded by sprouting many new stem suckers. The ability of Cercidiphyllum to resprout following stress (age, drought, fire, coppicing) is well known. Assuredly this aged survivor had once been a single stem, hundreds of years before even Wilson photographed it. Years later, Wilson recalled this visit to the tree in August of 1910, stating that it was actually the first time he ever saw the species ARNOLD ARBORETUM ARCHIVES Retracing Footsteps LIVING COLLECTIONS 7 Wilson photographed the leviathan Cercidiphyllum with three of his collectors on August 17, 1910, in what he described as \"open country.\" By 2017, the same tree and the forest around it had rebounded considerably, dwarfing NACPEC17 collectors (l to r) Kang Wang, the author, and Andrew Gapinski as they recreate Wilson's shot. ANDREW GAPINSKI 8 Arnoldia 75\/3 ? February 2018 Boston in early 1911. Our 2017 team was lucky, too, for this huge Cercidiphyllum bore fruits, which we collected under number NACPEC17-020 (we also collected fruits from other trees in the population under NACPEC17-021). And, luckily, none of us broke an appendage. Of Wilson's 1910 seed collection, but a single tree grew in the Arboretum. It was sited on Peters Hill, just to the south of the summit's presentday access road in an area then called \"rare trees of the Arboretum.\" Accession 7281*A was near an American beech (Fagus grandifolia, accession 22798*E) that remains to this day. This Cercidiphyllum had survived the brutally cold winters in the earlyto mid-1930s, proving its hardiness. However, by 1946, for an undocumented reason, it was dead. The tree had previously yielded enough budwood to produce five additional accessions, and a grafted plant (accession 133-41*B), the last of its lineage, grew in the Cercidiphyllum collection off Meadow Road. It was alive in 1948, but died shortly thereafter when its base snapped off, likely due to graft-incompatibility. In early November of 2017, just weeks after I returned from China and the Arboretum propagator sowed the seed, the first seedlings of both The short-shoot spurs of the large Cercidiphyllum japonicum bore rounded of our 2017 collections germinated to cordate leaves, waxy on the undersides, and small green fruits (follicles), with abandon. In a few years, it will which the team gleefully harvested. be a privilege and a joy to reintroin fruit. He added parenthetically, \"Later I colduce an exact Wilson acquisition to the collected ripe seeds, and this tree is now growing lections. While no doubt some trees will go in the Arnold Arboretum, where it promises along Meadow Road and elsewhere, I think one to be quite hardy.\" Technically, it would have should return to the spot near the Peters Hill been Wilson's collecting team that acquired the Summit, for old time's sake. seeds and fruiting vouchers that October, for on Literature cited September 3rd, 1910, a few weeks after initially Wilson, E. H. 1929. China, Mother of Gardens. 408 pp. seeing this Cercidiphyllum, he was caught in a The Stratford Company, Boston. landslide that busted his leg in multiple places. Luckily, his steadfast companions carried him to Chengdu where he spent the remainder of Michael S. Dosmann is the Keeper of the Living Collections at the Arnold Arboretum. the autumn recuperating, before returning to "},{"has_event_date":0,"type":"arnoldia","title":"Betula pumila: A Dwarf Among Giants","article_sequence":3,"start_page":9,"end_page":12,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25632","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15ea76b.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Enzenbacher, Tiffany","article_content":"LIVING COLLECTIONS 9 Betula pumila: A Dwarf Among Giants JOSHUA MAYER Tiffany Enzenbacher W hen one conjures an image of a birch (Betula spp.), typically a majestic tall tree with graceful architecture comes to mind--certainly not a low-growing, widespreading shrub. But, the small-statured bog or low birch (Betula pumila) is exactly what the 2017 Wisconsin Expedition (WIE) team, Manager of Plant Records Kyle Port and I, pursued from 23 August to 3 September of 2017. Since I began employment at the Arnold Arboretum three years ago, I have viewed our low birch accessions on an almost daily basis. Due to their short and scrubby growth habit, the specimens grow at the Arboretum alongside other dwarves: the plants of the Bonsai and Penjing Collection. Even as a caretaker of our dwarf potted plants, never did I imagine that I would be seeking B. pumila. The Arboretum has record of receiving 11 Betula pumila accessions prior to 2017. Founding Director Charles Sprague Sargent obtained the inaugural accession in 1876 from Mount Mansfield, Vermont just four years after the Arboretum's inception. Presently however, just two living accessions exist, the first comprising the two plants (800-93*A and B) growing next to the Bonsai and Penjing Pavilion. Jack H. Alexander III, former Arnold Arboretum Plant Propagator at the Dana Greenhouses (1976 to 2016), collected those seed-bearing catkins in Gros Morne National Park in Newfoundland, Canada in the fall of 1993. Our second accession (660-2016) is still in production at the Dana Greenhouses. We received small plants from the National Plant Germplasm System of the USDA-ARS (United States Department of Agriculture ? Agricultural Research Service) in 2016, with the seed originally harvested from Bremer County, Iowa. As the Manager of Plant Production, I obviously have regular check-ins with these seedlings. When caught at the right moment in early spring, the pistillate (female) catkins of low birch can be striking. KYLE PORT 10 Arnoldia 75\/3 ? February 2018 Coordinates from a 1958 herbarium voucher helped the collecting team locate this low birch (Betula pumila) population, still thriving in the drainage ditch bog along Highway 54 in Jackson County, Wisconsin. The ripening pistillate (female) catkins of low birch are bright green, nestled amidst the rounded to ovate, toothed leaves. KATHLEEN DOOHER Low birch is the only shrub birch native to Wisconsin and is widespread throughout the United States, indigenous to the northern Midwest, West Coast, Northeast, as well as much of Canada. It occurs in a variety of wetlands, such as bogs (areas of soft, water-logged ground), fens (low lying, frequently flooded land), and swamps (wetlands dominated by woody plants) in calcium-rich regions. During our expedition, Kyle and I anticipated finding abundant plants because the majority of the Nature Conservancy preserves we planned to visit harbor these bodies of water. After several days of looking in these prime habitats, to my utter disappoint, not a single Betula pumila was found. Viewing the Arboretum's only ex situ accessions every day for years had made this the one target taxon that I truly desired to acquire. So on our expedition's fifth day, Kyle and I decided to try another approach and go on a plant hunt using coordinates we pulled from a 1958 herbarium voucher in the Wisconsin State Herbarium database. The point was off Highway 54 in Black River Falls, above the Wildcat Ridge State Natural Area. The record indicated that the bog was being drained, and that black chokeberry JOSHUA MAYER Betula pumila LIVING COLLECTIONS 11 The author counts and cleans the tiny seeds--winged nutlets--of Betula pumila before direct sowing, or placing them under cold stratification at 2.2?3.3?C (36?38?F) for 3 or 4 months. These three different treatments will evaluate which brings greatest germination success. (Aronia melanocarpa), eastern larch (Larix laricina; another WIE priority taxon), and black spruce (Picea mariana) were associated species. After navigating to our location using our GPS, Kyle and I parked the vehicle, walked a short distance from a turnout, and began to look around. We excitingly noted that black chokeberry was still abundant, and walking further down the highway's shoulder, we were elated as we saw the 1950's low birch population flourishing 60 years later. At last, my long sought-after shrub was right before me in its native environment, after being so elusive the entire expedition. As the shrub groupings were not accessible 12 Arnoldia 75\/3 ? February 2018 (1 ? inches) long by 3.2 centimeters (1 ? inches) wide, and oval or slightly orbicular (round) in shape. And luckily, the plants harbored persistent female catkins that contained small, 3.2 millimeter (1\/8 inch), winged nutlets. Kyle and I harvested as many catkins as possible from the low birches in the vicinity, placed them in a labeled cloth bag, and later that afternoon mailed them overnight to the Dana Greenhouses with other bounties accumulated over two days' time. Upon my return from the field, I insisted on cleaning and processing the seed myself. Later as I removed the chaff and counted thousands of nutlets, I thought it fitting that once again a team member from the Dana Greenhouses, keepers of the Bonsai and Penjing Collection, was the one to collect this unusual dwarf shrub in a tree genus. Tiffany Enzenbacher is the Manager of Plant Production at the Arnold Arboretum. TIFFANY ENZENBACHER from the road, we carefully made our way into the bog using sedge (Carex spp.) clumps as stepping stones. As we got closer, Kyle and I went over the identifying traits on our mental checklist to validate that they were indeed low birches. They stood approximately 2.5 meters (8 feet) tall, which is in the 0.9 to 2.7 meter (3 to 9 feet) range. This is a stark contrast to the 20-meter (65-feet) height attained by another native Wisconsin birch species, yellow birch (Betula alleghaniensis), which we collected the following day. The other features of these plants were the same as the very familiar Arboretum accessions and the description that Kyle and I had memorized. New twigs were characterized by smooth, reddish brown bark with speckled white lenticels (raised pores on stems that permit gas exchange between the environment and plant tissue). Dentate (bluntly or sharply toothed) leaves were small, 3.8 centimeters Manager of Plant Records Kyle Port collects new GPS coordinates of a low birch specimen. "},{"has_event_date":0,"type":"arnoldia","title":"Coastal Southeast Expedition 2017: How Habitat Type Guides Collecting","article_sequence":4,"start_page":13,"end_page":15,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25633","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15eab6f.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Zukswert, Jenna; Halloran, Sean","article_content":"LIVING COLLECTIONS 13 Coastal Southeast Expedition 2017: How Habitat Type Guides Collecting CAMERON STAND Sean Halloran and Jenna Zukswert The COSE team included (l to r): Cat Meholic (University of Delaware), Ethan Kauffman (Stoneleigh, Natural Lands Trust), Jenna Zukswert (Arnold Arboretum), Jessica Slade (Morris Arboretum), Sean Halloran (Arnold Arboretum), and Tom Clark (Polly Hill Arboretum and Mount Holyoke Botanic Garden). Not pictured are local collaborators Richard Porcher, Mike Ammons, Gary Kauffman, and Andy Walker, who led the team to wonderful wild places. T he Campaign for the Living Collections takes explorers to many diverse parts of the temperate world in search of target taxa. Specific knowledge of plant ecology and plant communities helps us determine where we can find these taxa, and therefore where we will travel. From October 12?19, 2017, the multi-institutional Coastal Southeast Expedition (COSE) brought us to the coastal plains of South and North Carolina. Traveling through national forests, private lands, and public parks, we saw many habitat types, including calcar- eous forests, maritime depressions, pocosins (wetland bogs), Carolina Bays, longleaf pine savannas, pond cypress swamps, and ecotones (transitional regions between these habitats). In covering several hundred miles of Atlantic coastal plain between these two states, we targeted and collected taxa successfully by understanding and interpreting the plant community associations inherent in these various habitat types. The classic paradigm of plant collecting involves gathering historical locations of desired SEAN HALLORAN 14 Arnoldia 75\/3 ? February 2018 We then traveled to the Sewee Shell Ring in Francis Marion, where an interpretive trail leads to a 4,000-yearold shell ring and an 800-year-old clamshell mound. These shell mound sites, called middens, were created by Native Americans discarding clam and oyster shells that have since broken down into soil that favors the growth of certain plant species. By seeking out a sandy shell-influenced soil type at a maritime forest edge, we found Opuntia humifusa (eastern prickly pear) and Tilia americana var. caroliniana (carolina basswood). Just inland The prickly pear cactus, Opuntia humifusa, grew in well-drained shell from these collections we found true mounds. This target species may wind up in the Arnold Arboretum's maritime forest, with heavier soils recently renovated Rockery. and accordingly an abundance of taxa, while leveraging local knowledge through stately Quercus virginiana (southern live oak) floristic experts in the targeted collection draped in Tillandsia usneoides (Spanish moss), areas. During COSE, we saw the importance of with wonderful twisted trunks overhanging plant collectors combining these classic ideolosalt marsh flats. gies with the knowledge of ecological fundaAnother habitat we encountered was the mentals such as floral associates, environmental longleaf pine savanna. Once covering more than factors, and lifecycle needs to identify and locate 60 million acres in the American southeast, our target taxa. these habitats, dominated by Pinus palustris Our trip began in a calcareous, or limestone, (longleaf pine), have shrunk to less than 3 milbluff forest in the Francis Marion National lion acres due to overharvesting and deforesForest in South Carolina, where we collected tation. Despite the overwhelming dominance Acer barbatum (southern sugar maple). By of P. palustris, these ecosystems are actually seeking out this rare forest type in South Caroamong the most diverse in North America, with lina, which is unique in its exposed rock outplant diversity levels in the understory among croppings and high-calcium soils, we found the highest outside of the tropics (Outcalt and the home of several species that are otherwise Sheffield, 1996). Many of the herbaceous sperare in this region. We collected seedlings of cies found in these habitats depend on condiA. barbatum, and found Sabal minor (dwarf tions created by the presence and dominance of palmetto), Ulmus rubra (slippery elm), Juglans P. palustris, or on frequent fires. Without fires nigra (black walnut), and various ferns. Just inhibiting woody plant expansion, the underdown the bluff was Wadboo Creek, a tributary story would not be nearly as diverse. of the Cooper River. In this wooded swamp, We first sought out this ecosystem type in we expected to find plants that would thrive South Carolina, and collected Pinus palustris in a similar soil type, but with much more cones in conservation lands at Brookgreen available water, such as Cornus foemina Gardens, led by their natural lands manager (swamp dogwood) and Ampelaster caroliniaMike Ammons. This pine savanna had dry, sandy soils, which don't occur frequently in the nus (climbing aster), the latter of which was coastal plain, and we expected to find P. palustris climbing other shrubs and trees on both sides trees in all stages of growth, given its tolerance of the creek. We were just in time to enjoy its to fire and need for sandy, well-draining soils. yellow-centered lavender flowers. SEAN HALLORAN CAT MEHOLIC Coastal Southeast Expedition LIVING COLLECTIONS 15 plants, we collected Persea palustris (swamp bay), Gordonia lasianthus (loblolly bay), and Zenobia pulverulenta (dusty zenobia). We learned from Andy and Gary how quickly these plants re-establish after fire: within two seasons, bur ned areas become nearly impassable to humans due in part to regenerative shrub growth, as well as the vining Smilax laurifolia (laurel greenbrier). Familiarity with the ecosysThe longleaf pine (Pinus palustris) savannah at Brookgreen Gardens in South tems of our target species is Carolina exhibited all stages of longleaf pine development from grass, bottlevaluable not only for finding and brush, and sapling stage (first branching), all the way up to mature trees with collecting them, but also for concharred trunks. The species can stay in its grass stage for many years, and then in a single year grow rapidly and tall enough to escape the effects of potential fire. sidering the next phases of life for our propagules. While the Arnold Arboretum landscape does not feature the same conditions that we encountered in the Carolinas, we can attempt to replicate some of their characteristics when siting the COSE collections as Arnold Arboretum accessions. Luckily, with each germplasm collection, we took copious notes describing the plant and the environment where we found it--this documentation is just as important as the germplasm itself. To give our collections the best Longleaf pine savannas in South Carolina support an array of fascinating chance to thrive in Boston, we understory species, including Asclepias humistrata (sandhill milkweed), with a can consider these data with other sprawling growth habit in sandy areas that makes it appear to grow sideways. criteria pertaining to the ArboreIn North Carolina, we encountered longleaf tum landscape, such as topography, soil condipine savannas in the Croatan National Forest. tions, water status, and even how the original Botanists Andy Walker and Gary Kauffman species associates perform here. introduced us to this habitat and pointed out Literature cited rare plant species, such as federally endangered Outcalt, Kenneth W. and Raymond M. Sheffield. 1996. Lysimachia asperulifolia (roughleaf yellow The longleaf pine forest: Trends and current loosestrife). These savannas were wetter than conditions. Resour. Bull. SRS?9. Asheville, NC: those we visited in South Carolina, and thus U.S. Department of Agriculture, Forest Service, had more species tolerant of wet conditions. Southern Research Station. 28 pp. In small, wet depressions, for example, we found several Sarracenia (pitcher plant) Sean Halloran is the Plant Propagator and Jenna species as well as Dionaea muscipula (Venus Zukswert was a Living Collections Fellow (2016?2017) at the Arnold Arboretum. fly trap). In addition to these carnivorous "},{"has_event_date":0,"type":"arnoldia","title":"Pieris phillyreifolia: The Opportunistic Climbing Fetterbush","article_sequence":5,"start_page":16,"end_page":18,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25637","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15eb36b.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Halloran, Sean","article_content":"16 Arnoldia 75\/3 ? February 2018 IMAGE COURTESY OF BIODIVERSITY HERITAGE LIBRARY Pieris phillyreifolia: The Opportunistic Climbing Fetterbush Sean Halloran A t its core The Campaign for the Living Collections is a strategic endeavor involving years of planning and ten years of execution. Each expedition is organized thoroughly in advance, and anchored by specific target species or desiderata. However, no amount of planning can account for all of the factors at play when collecting plants in the wild, and as such non-target species are often collected opportunistically. Opportunistic collections can include biological outliers, like Pieris phillyreifolia (climbing fetterbush), the only Ericaceous (belonging to the heather family) plant native to North America that is also a woody vine or liana. After seeing this unique Pieris species, I could not resist the chance to collect it for the Arnold Arboretum, as a participant in 2017's Coastal Southeast Expedition (COSE). Pieris comprises seven known species. The two most utilized as ornamentals are the North American native Pieris floribunda (mountain andromeda) and Pieris japonica (Japanese andromeda). During our trip, we sought out the lesser-known P. phillyreifolia, found only in South Carolina, Georgia, Florida, and coastal areas of Mississippi and Alabama. Walter Judd (1982) wrote about this species in detail, but W.J. Hooker first described it in 1837, initially placing it in the genus Andromeda (Lemon and Voegeli, 1962). An 1837 illustration of Pieris phillyreifolia, appearing in Volume 2 of W. J. Hooker's Icones plantarum, where he originally placed it in the genus Andromeda. The specific epithet (phillyreifolia) reflects the resemblance of its leaves to those in the Mediterranean genus Phillyrea, in the olive family (Oleaceae). Pieris phillyreifolia LIVING COLLECTIONS 17 highway several times, Ethan got out of the car and motioned for us to join him in the woods: he believed he found the spot and the particular Taxodium ascendens (pond cypress) we were seeking. We all got out and trudged through the swampy borderlands of the highway to see if he was correct. He was! We found lianas just 9 meters (10 yards) from the highway. It is often unique plant form and function that interest people the most, and what makes the story of this species so fascinating is not just where and how we found, but what it grew on, and the mechanics of that growth. The common name, climbing fetterbush, is apt as we found it climbing on Taxodium ascendens in a cypress swamp. The root system was nestled in the buttressing roots of the cypress, and the stem traveled up the host tree underneath the bark, emerging from vertical cracks every 1 to 2 meters (3.3 to 6.6 feet) as aerial SEAN HALLORAN While exploring Francis Marion National Forest in South Carolina, we crossed paths with Clemson University's Patrick McMillan leading a group tour of the natural history of the area. Ethan Kauffman of Stoneleigh Gardens (one of the COSE participants) was keen to collect the unusual climbing fetterbush, and mentioned this to Patrick, who then produced a hand-drawn map of a nearby location of Pieris phillyreifolia for the team. Armed with this treasure map, our group eagerly set out in the National Forest along US Route 17 to find it. Our collecting routine was a bit comical: we would drive along the highway and then every few minutes Ethan would jump out and run into the woods, only to emerge seconds later to let us know we were not there yet. This occurred several times and admittedly, I was growing skeptical that we would find ever this population. Finally, after turning around on the Climbing fetterbush rhizomes grow underneath fissures in the fibrous pond cypress bark, emerging periodically to produce leafy green branches. shoots that extended outward up to half a meter (20 inches) from the tree. The plants we found were robust climbers reaching at least 6.1 meters (20 feet) up the Taxodium. On our trip last fall, we only observed this plant climbing T. ascendens, however it also grows on Chamaecyparis thyoides (Atlantic white cedar), Cyrilla racemiflora (swamp cyrilla), Pinus elliottii (slash pine), and other downed trees and soil mounds, most likely to avoid standing water (Lemon and Voegeli 1962; Judd 1982). As a plant propagator, it is invaluable to understand a plant's life cycle to grow it successfully in a controlled environment. For many taxa there are established protocols for germinating seeds, rooting cuttings, or grafting budwood. However, this is not the case for Pieris phillyreifolia. When in doubt we often look to established protocols for related species, so I decided to treat the seed via cold stratification for 90 days at 4 degrees Celsius (39 degrees Fahrenheit), which our propagation records indicate work for other species of Pieris. My first-hand knowledge of climbing fetterbush's natural environment will The white, bell-like flowers of Pieris phillyreifolia are borne in short, axillary racemes. also be put to good use as we figure Literature cited out where to cultivate this unique Southeastern U.S. native once we have successfully propaHooker, W.J., and J.D. Hooker. 1872. Icones Plantarum, Or Figures, with Brief Descriptive Characters gated it. Given where we found it growing, I recand Remarks, of New Or Rare Plants, Selected ommend we site this plant in a wet depression from the Author's Herbarium (Vol. 12). London: under heavy deciduous shade where it can grow Longman. up a trunk or trellis. Given its native range, we Judd, W.S. 1982. A taxonomic revision of Pieris (Ericaceae). should protect it from harsh winters by estabJournal of the Arnold Arboretum 63:103?144. lishing it in a warmer microclimate within the Lemon, P.C., and J.M. Voegeli. 1962. Anatomy and ecology Arboretum. Observing a plant's unique form of Pieris phillyreifolia (Hook.) DC. Bulletin of in the wild, and researching its fascinating histhe Torrey Botanical Club 89:303?311. tory upon return, deepens the appreciation for all propagules collected afield. The climbing Sean Halloran is the Plant Propagator at the fetterbush is no exception. Arnold Arboretum. CHRIS EVANS, UNIVERSITY OF ILLINOIS, BUGWOOD.ORG 18 Arnoldia 75\/3 ? February 2018 "},{"has_event_date":0,"type":"arnoldia","title":"Three Times a Collection: The Quest for Moonseed","article_sequence":6,"start_page":19,"end_page":21,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25639","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15ebb28.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Woodruff, Kea","article_content":"LIVING COLLECTIONS 19 Three Times a Collection: The Quest for Moonseed Kea Woodruff F or members of Arnold Arboretum plant expedition teams, the target taxa list is their guiding document: What are we looking for and where will we find it? Being interested in all things botanical, each collector is also invariably curious about which plants are on the other teams' target lists. When these species are duplicated on a few different teams' lists, a friendly competition arises that also unites them over a shared mission. As a firsttime collector accompanying Robert Dowell on the Arkansas-Missouri Expedition (ARMOE), I was eager to test my field identification skills and compare expedition stories with the other two domestic teams in 2017. If the others were picking beechnuts in the Carolinas, I wanted to find them in Arkansas. If they were digging up red maple in Illinois, I wanted to find it in Missouri. There were several species targeted by multiple teams, but only one was successfully collected across all three domestic expeditions--Menispermum canadense (Canada moonseed). Menispermum is a temperate genus in a primarily tropical family, and contains just two species: Menispermum dauricum from northeastern Asia, and M. canadense, found throughout much of eastern North America. The Arnold Arboretum has only one accession of this woody vine, obtained in 1994 from a ? ? ? Common moonseed occurs throughout much of eastern North America, and the collections (?) in 2017 represent the north-northwestern, southwestern, and southeastern portions of its range within the United States. Dark green portions of the map illustrate states or provinces of species' nativity, with specific counties of occurrence shown in light green. Counties where the species is rare are shown in yellow, while counties with historic concurrences are in orange. Map modified from: Kartesz, J.T., The Biota of North America Program (BONAP). 2017. Taxonomic Data Center. (http:\/\/www.bonap.net\/tdc). Chapel Hill, N.C. JAN DE LAET, PLANTSYSTEMATICS.ORG 20 Arnoldia 75\/3 ? February 2018 The broad, palmately lobed leaves of Menispermum canadense are up to 24 centimeters (9.5 inches) wide and long. Small white flowers develop into clusters of round, green and then purple fruits (drupes) when ripe. cultivated source. The absence of wild material prompted its inclusion on the target list. Not only did our successes fulfill the Campaign goal of bringing in material of wild origin, but because they came from diverse geographic regions, they collectively improve the robustness of the Arboretum's holdings. Growing multiple accessions of a species from across its native geographic range increases the overall genetic diversity represented in the Living Collections. These different individuals are likely to exhibit a broader range of species' traits and adaptations, leading to a richer resource for researchers and visitors alike. And, generally speaking, growing multiple accessions of different provenances increases the odds of success (although this species is rather easy to cultivate). Canada moonseed is not a showy vine-- a former Arboretum researcher once called it \"charming but overlooked\" (Young, 2014). Plants can climb up to 5 meters (16.5 feet) tall, and can be mistaken for various species of Vitis, or wild grape. Unlike wild grape, however, all parts of the Canada moonseed are toxic. For the plant collector working in the field, diagnostic features are crucial in identifying the correct plant. A lack of tendrils--the threadlike tissue vines use to coil or spiral around supports-- in Menispermum is a critical feature that can help distinguish it from its lookalikes. Another important diagnostic feature is its distinctive, crescent moon-shaped seed, from whence it takes its common name. Seed is a fickle commodity. For the plant collector, locating a target species and then finding it barren of fruit can be a disappointing experience. Fortunately, many species can be collected through other means. Seedlings or young plants are an alternative if granted permission and the local plant community will not be negatively affected. Collectors will often dig up seedlings early in an expedition, hedging their bets in case they never find seed later on. Such was the case in 2017. In Missouri, my collecting partner Robert Dowell identified a SEAN HALLORAN Menispermum canadense LIVING COLLECTIONS 21 The moon-shaped seed collected in Wisconsin (Accession 285-2017) germinated readily upon sowing. creek bed in Mark Twain National Forest as a likely habitat for Menispermum canadense. Looking in the understory beneath huge sycamore (Platanus occidentalis) trees, we found many vine seedlings, including Smilax spp. (cat briar), Vitis spp., and poison ivy (Toxicodendron radicans). It took close (and careful!) inspection to positively identify M. canadense among the others. Although seedlings were abundant, the lack of mature vines meant finding seed would be improbable, so we dug them up. The Wisconsin-Illinois team collected earlier in the fall, and found seed only after first collecting seedlings. Their stories kept us hopeful that the experience would be duplicated for us. And, sure enough, later that same day, while creeping slowly along the road, we found M. canadense plants bearing fruits which we harvested with much jubilation. The Coastal Southeast team eventually collected their seedlings in North Carolina, too. When we all got back and compared notes, there was a great sense of shared excitement that multiple teams collected this species, yielding six new accessions. All three teams found Menispermum canadense in shady understories, very near a creek, and in the same habitat as poison ivy. Once old enough to be planted out, these additions will provide an interesting native species to the shrub and vine collection. Easy to grow, this vine is generally free of insect predators and disease, and tolerates a range of conditions from full sun to full shade when in cultivation. This broad adaptability increases the likelihood that individuals from all three collection sites will thrive in Boston, giving Arboretum staff and future visitors a chance to observe and study potential differences in habit and performance among them. Bibliography Ortiz, R.D.C., Kellogg, E.A., and Van Der Werff, H. 2007. Molecular phylogeny of the moonseed family (Menispermaceae): Implications for morphological diversification. American Journal of Botany 94: 1425?1438. Young, S.T. 2014. Untangling the ways of lianas. Silva (The Newsletter of the Arnold Arboretum) 14:10. Kea Woodruff is the Plant Growth Facilities Manager at the Weld Hill Research Building. "},{"has_event_date":0,"type":"arnoldia","title":"Going Clonal: Beyond Seed Collecting","article_sequence":7,"start_page":22,"end_page":23,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25634","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15eaf28.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Dowell, Robert","article_content":"KYLE PORT 22 Arnoldia 75\/3 ? February 2018 Going Clonal: Beyond Seed Collecting Robert Dowell S White, pea-like flowers of Cladrastis kentuckea are abundantly borne in long racemes, as in this old tree in the Arboretum's collection (accession 16370*A). Cladrastis kentuckea occurs in scattered, disjunct populations throughout the south-central United States. The large expanse in the westernmost part of the species range served as the source of the 2017 collection. Modified from Little, E. L. Jr. (1999). Atlas of United States Trees. U.S. Geological Survey. ROBERT DOWELL eed is the most important and most valuable propagation material an expedition targets. A handful of seed can offer genetically diverse, and logistically easy, material to procure and grow for the Arboretum's collections. Yet some target taxa present unique difficulties for collectors searching for seed. One species that exemplifies this is Cladrastis kentukea (American yellowwood). As part of the Campaign, American yellowwood is a target. This species is unique as the only member of its genus native to North America--all others occur in Eastern Asia. Furthermore, not only is it disjunct from its Asian relatives, but its North American populations are scattered in distribution. Of the 13 living accessions in the Arboretum landscape, only one (accession 51-87) has known wild origins, collected in 1986 by Rob Nicholson in Tennessee during the Southeastern States Expedition. Thus, to broaden the species' genetic diversity in cultivation in the Arboretum, we selected its westernmost range to source additional wild material. During the September 22 to 30, 2017 Arkansas-Missouri Expedition to the Ozarks (ARMOE), Kea Woodruff and I pursued Cladrastis kentukea. This tree can be a hard target to hit and offers several lessons for the plant collector. The species is nowhere abundant in its range, so pinpointing it can be an exercise in frustration. Thus, a previous collector's notes on the locations of existing populations, as well as observations of population health and size, can greatly aid a future collector's hunt. Luckily, we tracked C. kentukea in Arkansas due to the insights of a previous collector, Jeffrey Carstens, of USDA-Agricultural Research Service's North Central Regional Plant Introduction Station in Ames, Iowa. When collectors do find Cladrastis kentukea, they often find a small population of individuals producing few if any viable seeds. Even if a Root cuttings of American yellowwood, buried horizontally in a mixture of peat, perlite, and pinebark. CloningLIVING COLLECTIONS 23 collecting team is lucky enough to beat these odds and find a healthy population with high seed set, they must do so in August. The species disperses its seed before many others do--and before most collecting expeditions occur. These frustrations have played out for past Arboretum collecting efforts. In 1986, Rob Nicholson procured seven seedlings (four of which remain in the collections), because no seed was available in October. During the Southern Appalachian Expedition (SAPPE) of late September 2016, I observed the species in both North Carolina and Georgia, yet each small population also lacked seed. Thus, we were not hopeful to find seed-bearing American yellowwood on our trip. Luckily, Cladrastis kentukea illustrates an important yet less often approach for collectors: gather vegetative or clonal material instead of seeds. Depending on the season and the species, this could include leafy cuttings, dormant stem scions, or root cuttings. However, like a number of other woody genera in the legume family (Fabaceae), the only viable method for Cladrastis is to collect root cuttings, as enumerated by Peter Del Tredici (1995). With asexual or clonal propagation, the genetic diversity in any given collection is significantly lower than if you collect seed. A handful of sexually derived seed from one tree is a much greater pool of genetic diversity than a handful of cuttings from the same tree (which would yield identical clones). For this reason, when taking cuttings one should sample as many individual trees in a population as possible to maximize genetic capture. Armed with this knowledge of root cutting possibilities, and location data from Jeffrey Carstens, we set our sights on an American yellowwood population at the Long Pool Recreational Area, in the Ozark National Forest. This population is located upslope--and on a very steep slope at that--from a trail running adjacent Big Piney Creek. The dry understory was noticeably rocky, with the ever-present limestone bedrock well known in Arkansas. The population we encountered was small, about 20 mostly juvenile trees; many exhibited significant dieback. The largest tree, approximately 9.1 meters (30 feet) tall and with a diameter at breast height of about 30.5 centimeters (12 inches), was the victim of recent storm damage. The tree's main leader had snapped off, and was on the ground like a decaying skeleton. As we expected, there was no seed, but we were able to gather one to two root cuttings each from seven juveniles (cuttings from young trees tend to root more easily). After excavating a bit of the soil, we collected cuttings from as near the root crown or base of the tree as possible, being cautious to do as little harm as possible. This region also lies within the \"cone of juvenility\" that propagators know maximizes their chances for successful rooting. The cuttings themselves are generally 7.6 to 15.2 centimeters (3 to 6 inches) long and about 1.3 centimeters (0.5 inches) wide. When collecting them, the proximal end (the end closest to the trunk) is cut straight across whereas the opposite or distal end is cut at a slant. This allows the propagator to later identify the correct polarity or orientation of the cuttings if they insert them vertically in flat filled with growing media (proximal end up). However, typically cuttings are placed in the flat horizontally. Root cuttings generally do not require any hormone treatment (unlike stem cuttings) and are simply placed in a warm greenhouse environment to induce rooting. After considering the special circumstances in which to use them, asexual propagation techniques serve as additional tools in the toolbox that collectors can rely on to enhance their expedition success. Of the 68 taxa (104 accessions) collected on the three 2017 domestic expeditions, 11 taxa (14 accessions) represented collections of clonal material. Some of these were the more traditional leafy cuttings or rooted divisions, but others like the American yellowwood were of root cuttings. For example, Ulmus spp. (elm) disperse their seeds very early in the season, so Kea and I collected root cuttings of Ulmus alata (winged elm) and Ulmus rubra (slippery elm) on our trip. These elms were already sprouting shoots by early November 2017. Bibliography Del Tredici, Peter. 1995. Shoots from roots: A horticultural review. Amoldia 55(3): 11?19. Robert Dowell was a Living Collections Fellow at the Arnold Arboretum (2016?2017). "},{"has_event_date":0,"type":"arnoldia","title":"Recalling Plums from the Wild","article_sequence":8,"start_page":24,"end_page":34,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25638","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15eb76f.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"Recalling Plums from the Wild Jonathan Damery I BIODIVERSITY HERITAGE LIBRARY n 1811, a fur trader named George Sibley led a small team on a search for a storied salt mountain in the northern prairies of presentday Oklahoma. Sibley found the location in late June, although the salt did not mound; rather it formed a shimmering plain that stretched over dozens of snow-white miles. Bison peppered the expanse. On sandy hills rimming the salt plain, the team found shrubs, scantly waist high, that were loaded abundantly with small ripe plums. Sibley plucked these eagerly; they were, he said, \"the most delicious plums I ever tasted.\"1 Small native plums can be found across much of North America. The most widespread species, Prunus americana, ranges from New England to the Rockies, and it has garnered common names fitting for this range: the American plum or, more generally, the wild plum. Taxonomists disagree on the number of native plum species, but the Flora of North America includes thirteen, nine of which inhabit the central part of the continent, west of the Mississippi (see pages 32 and 33 for a gallery). Sibley probably waxed about the Oklahoma plum (Prunus gracilis), given the early fruiting time and sandy habitat, but despite such arduous praise (\"the most delicious\"), native plums seldom appear on grocery shelves or beneath farmers market tents. That has not always been the case. Luther Burbank, the famous horticultural polymath who began breeding plants at his home in Santa Rosa, California, in 1875, asserted that there were three important periods for plum cultivation: \"the wild era,\" \"the backyard era,\" and \"the railroad era.\"2 The disappearance of native plums occurred along this historic trajectory, at a collision between technology and taste. Selecting Plums from the \"Wild Era\" This image of Orville Lord at the age of 77 appeared in the June, 1902 issue of The Minnesota Horticulturist. The annotation below is in his own handwriting. When Charles Sprague Sargent, the first director of the Arnold Arboretum, wrote about plums in his fourth volume of the Silva of North America, published 1892, he noted that the fruits of several native species were common in markets, particularly in inland cities like St. Louis, where foraged plums were sold both fresh and jellied. Pomologists in Iowa, Minnesota, Wisconsin, Texas, and elsewhere were also selecting cultivated varieties or cultivars (at the time referred to simply as varieties) with larger fruits, thinner skins, and freestone pits. \"Selected varieties sometimes produce PETER ASSMANN Native Plums 25 The yellow to red fruits (drupes) of the Oklahoma plum, Prunus gracilis, range from 1 to 2 centimeters (0.4 to 0.8 inches) in size. excellent fruit,\" Sargent wrote, \"and have been largely cultivated, in the western states especially, for many years.\"3 While the Arboretum did not grow any native plum cultivars at the time, Sargent would have received insights about breeding and selection efforts from the pages of Garden and Forest, the horticultural magazine he began editing in 1888. In an article from 1891, Emmett Stull Goff, the first professor of horticulture at the University of Wisconsin, recounted a field trip to an orchard in southeastern Minnesota, along the banks of the Mississippi. The owner, Orville Lord, had gained regional acclaim for native plum cultivation, shipping fruits as far as New Jersey. Lord had introduced a cultivar of American plum that he named after a nearby creek, `Rollingstone.' Goff compared the fruit of `Rollingstone' favorably to `Green Gage,' a popular cultivar of domestic plum (Prunus domestica), which, even today, occasionally appears in grocery bins. Fruits of `Rollingstone' and `Green Gage' were about the same size, Goff reported, and although the skin of `Rollingstone' was thicker, he conceded that the native was nonetheless \"delicious.\"4 Horticultural experimentation with native plums occurred for practical reasons. While domesticated plums had been imported from Europe, where plum consumption has occurred since Roman times, if not earlier, 5 pests and diseases proved persistent obstacles for orchardists in the central United States. Black knot, the fungal disease that forms aptly named lumps on plum branches, was one of the chief problems, as was the plum curculio, a weevil that feeds on flowers and greening buds and eventually young fruit. Although the native species were not completely immune from these problems, they fared significantly better.6 Orchardists in northeastern states had better luck with classic European cultivars, so interest in native cultivation remained primarily within the Mississippi watershed. Liberty Hyde Bailey, at Cornell University, listed 140 cultivars of native plums in his 1892 publication, The Cultivated Native Plums and Cherries, but it is clear that he had not grown many himself.7 Rather his curiosity had been aroused by the vexing taxonomy of the species. Taken as a whole, Bailey thought native plums represented a classic instance of \"contemporary evolution,\" given the high-degree of hybridization and morphological plasticity. His descriptive list of cultivars included fruit reviews, flowering times, and provenance narratives, but even this seemed to straddle a dual function: a horticultural guide for would-be orchardists, coupled with an attempt to systematically describe the range and variability of particular species. Cultivated varieties provided Bailey with useful taxonomic information because, according to his estimation, more than half were wild-collected favorites, imported directly from the hedgerow to the orchard. Significantly, none of the 45 cultivars of American plum on Bailey's list originated from wild locations east of Illinois, despite a Hedrick, in The Plum's of New York, considered `De Soto' \"first place species range that extends all the among the American plums\" for its productivity and ability to withstand shipping. The cultivar was discovered on the banks of the Mississippi River way to New England. 8 Most came near De Soto, Wisconsin in either 1853 or 1854. from Minnesota, Wisconsin, Iowa, and occasionally Missouri. One of these wild as have other tribes across the continent.9 Lord selections was Lord's `Rollingstone,' which was instantly enamored with the large, sweet he first encountered in 1852, the same year fruit, although he would not introduce `Rollinghe arrived in southern Minnesota and setstone' to market for about three decades, when tled among mounded Mississippi bluffs. The his attention, in older age, shifted evermore original shrub was growing on the edge of a towards horticulture.10 seasonal settlement used by the Mdewakanton-- In Minnesota City, Lord attempted to grow a subgroup of the Dakota--who may have every cold-hard variety, reporting in 1903 that intentionally selected and planted it near their he was cultivating more than one hundred disencampment. Certainly, the Dakota have long tinct selections.11 The most extensive trial, valued native plums, both fresh and preserved, however, likely belonged to Jonathan Kerr, a BIODIVERSITY HERITAGE LIBRARY 26 Arnoldia 75\/3 ? February 2018 Native Plums 27 nurseryman in Denton, Maryland. Bailey spent a considerable amount of time at Kerr's Eastern Shore Nurseries, researching varieties before he published his plum report. Unlike Lord, who primarily raised native plums for commercial fruit production and experimentation, Kerr intended to supply homeowners and orchards with plant material. In an 1895 catalog, Kerr announced that orchardists near Baltimore and Philadelphia had sold native plums for up to four times as much as the domestic plums. \"They pay better,--the pay oftener, than any other tree fruit,\" the catalog promised.12 The following year, Kerr reported that the nursery was growing more than 250 varieties.13 Over the next decade, that figure would double.14 Tracing Plums into Backyards and Orchards It might be careless to assume a proliferation of cultivar names implies a corresponding proliferation of cultivated acres. William Wight, a botanist for the U.S. Department of Agriculture, noted the cavalier nature of many of the horticultural selections. In a taxonomic report on the species, published in 1915, Wight estimated that more than 800 selections had been named, and some of these, he suggested, were \"no better, doubtless in some cases not so good, as those found in a wild state.\"15 Even so, the native plum industry was far from mere nursery hucksterism. According to the U.S. Census of Agriculture--a report, established in 1840, tabulating everything from acres of barley to pounds of butter and fertilizer expenditures--plum production exploded throughout the central United States during the final decade of the nineteenth century. Iowa emerged as a leader, with more than 1.3 million plum trees under cultivation, almost five times the amount reported a decade before. Illinois and Missouri increased at similar rates, amounting to more than a half million trees for Illinois and three-quarters of a million for Missouri.16 While the census did not delineate between species of plum, the authors noted that \"Chickasaw and allied varieties\" predominated in the \"Mississippi Valley.\" 17 This assessment echoed recommendations at state horticultural society meetings throughout the region, where native cultivars were always the most praised and discussed. \"Our natives are the only sure foundation for commercial plum orchards in Iowa,\" an orchardist from Cedar Rapids, Iowa announced at one of these characteristic local meetings in 1896.18 Return to the Hedgerows While the U.S. Census of Agriculture traced the rise of the plum, it also recorded the subsequent bust. Almost a million fruit-bearing plums disappeared from Iowa over the first two decades of the twentieth century. Illinois production was halved over the same period, and Missouri plums also dwindled to almost half (see data, next page). If the rising number of cultivated names paralleled an explosion of cultivated acres, then the same trend seemingly held true as production of nursery stock dwindled. The catalog for Kerr's Eastern Shore Nurseries listed only nine native cultivars in 1914, along with two hybrids that claimed native parentage, far from nearly two hundred selections previously advertised. Now, more than a century after the native plum boom, most selections have vanished from markets and from cultivation at large. The U.S. Department of Agriculture's National Plant Germplasm System maintains repositories to conserve genetic diversity for future crop breeding. The plum collection is located in Davis, California and offers the most probable location to encounter an assortment of historic native plums. Yet compared to the number of named varieties offered for sale in Kerr's 1896 nursery catalog, the diversity is slim. The collection includes thirteen accessions of American plum, a majority representing wild provenances. Of five with cultivar names, only `Wolf' was included on Kerr's list of more than 110 American plum selections,19 although another (`Anderson') was also a nineteenth-century selection. Both, incidentally, were wild-collected from Iowa. My recent search for Orville Lord's orchard in Minnesota City, Minnesota, turned up no fruit, except for several wild American plums growing near a boat launch about a mile away from his property. The center of his land is now 28 Arnoldia 75\/3 ? February 2018 Midwestern Plum Cultivation American plum production trends resemble a slow partner dance, as orchards in one region expand concurrently with reductions elsewhere. Native plums dominated cultivation in the Mississippi Valley--including states depicted here--at the dawn of the twentieth century, but as the national market for California hybrids grew, Midwestern production crashed. California growers eventually outcompeted themselves, creating an oversupply revealed with plummeting Depression-era fruit prices.1 In the late 1950s, demand for canned plums encouraged additional Michigan production, but as consumer taste shifted towards fresh fruit, swelling California cultivation was once again cited as critical competition.2 Data extracted from the U.S. Census of Agriculture represent the number of fruit-bearing plum trees reported from 1890, the first census to include plum data, through 1997. Subsequent reports have noted acreage rather than tree counts. N U M B E R O F F R U I T - B E A R I N G P LU M T R E E S 1890 1900 1910 CALIFORNIA 1,509,833 9,823,713 7,168,705 ILLINOIS 104,111 572,774 600,087 INDIANA 146,378 723,815 566,988 IOWA 260,600 1,302,217 1,155,041 KANSAS 410,426 852,702 624,648 MICHIGAN 168,318 1,378,952 464,917 MINNESOTA 47,458 191,313 233,736 MISSOURI 152,688 745,187 917,851 NEBRASKA 227,129 542,450 351,321 NORTH DAKOTA 681 4,745 19,147 OHIO 145,832 892,441 1,001,734 SOUTH DAKOTA 42,797 123,175 268,268 WISCONSIN 18,451 94,338 105,909 1Matthews, G. 1985. The apricot war: A study of the changing fruit industry during the 1930s. Agricultural History. 59(1): 25?39. 2Ricks, D. J. 1983. The Michigan and U.S. purple plum industry--Trends and changing marketing patterns. Michigan State University Agricultural Economics Staff Paper. 83(56): 1?46. *(D) signifies \"withheld data\" a small subdivision, and a separate orchard of his in the Mississippi bottomlands was flooded when the Army Corps of Engineers built a lock and dam in 1935. Plums of the \"Railroad Era\" The disappearance of native plum cultivars can be partly explained by matters of taste. Anyone who has foraged one of these plums will likely describe the astringency of the skin, even while savoring the sweetness of the flesh inside. This characteristic is generally true of all native species. Ulysses Hedrick, who joined Bailey at Cornell, wrote a 1911 monograph on plums. He noted that while the American plum had been introduced to Europe in the mid-eighteenth century, if not before, the species was always considered a flowering ornamental in European gardens, not an orchard plant. \"The Old World plums are so superior, speaking generally, in size, appearance, and flavor, the qualities which appeal to those who eat plums, that the native varieties stand small chance for popular favor,\" he wrote.20 Still, work of orchardists like Orville Lord might have continued in the central United States if not for technological innovations. In 1887, Lord imagined a native plum industry that could surpass the $2.5 million market for imported plums and prunes. \"Does this sound visionary,\" he exclaimed at a meeting of Minnesota horticulturists. \"I may ask who would Native Plums 29 NUMBER OF FRUIT-BEARING PLUM TREES 1920 CA 1930 1940 1950 1969 1978 8,971,175 9,809,553 9,522,743 1987 1997 8,768,436 16,668,590 273,554 160,494 98,382 62,148 5,765 1,659 16,511 1,623 698 214,202 117,713 59,155 42,181 7,467 4,197 4,369 1,990 603 313,769 290,613 177,375 118,018 11,076 4,950 824 487 426 143,473 139,590 46,665 32,434 4,684 1,122 1,241 302 544 377,123 312,899 204,022 264,976 237,325 579,239 480,651 315,164 131,085 193,668 224,974 160,947 130,449 10,337 2,022 2,243 1,299 2,501 528,649 317,598 239,804 121,032 15,426 1,325 4,394 2,910 544 86,183 118,133 36,173 33,574 4,812 209 441 274 (D)* 41,254 47,423 14,657 48,989 19,573 1,779 1,340 (D)* (D)* 459,265 323,731 187,548 130,587 44,651 32,518 17,151 10,182 5,165 117,677 100,185 21,137 63,222 4,302 802 769 266 628 117,844 126,538 102,891 83,242 10,522 2,913 4,345 2,838 878 IL IN IA KS MI MN MO NE ND 12,915,324 10,285,039 1959 13,866,499 15,909,878 OH SD WI have dared to predict, thirty years ago, the small fruit business of ... Chicago, Minneapolis or St. Paul. Then, a carload would have supplied the market of either place. Now, thousands of bushels are daily marketed in their season.\"21 Lord was optimistic about the voracious appetite of a booming city. Over the same thirty-year period, Chicago grew from a city of about three hundred thousand to a metropolis exceeding one million, but Lord missed the implications of the same appetite. If Chicagoans could each consume one quart of strawberries--berries that were not grown in the city, but rather, were grown in the hinterlands and shipped inward via rail--then the same transport innovations could undercut the need for locally grown native fruits. Over the decades that Lord cultivated increasing acres of native plums, railroads had connected the coasts. The first transcontinental passage occurred in 1869, and, in 1890, a California fruit shipper, Edwin Earl, devised a railcar suited for long coast-tocoast shipments. Along the way, pantries and iceboxes in Chicago and beyond became less beholden to horticultural limits for the region. Luther Burbank alluded to these innovations when he posited the \"railroad era\" as the ultimate stage for plum production. \"The railroad became a factor in plum improvement by bringing millions of plum-hungry easterners within reach--by affording quick and economical shipping facilities where there had been no Horticulturist Luther Burbank, flanked by Thomas Edison (left) and Henry Ford (right), in this image that appeared in Burbank's 1916 seed catalog Twentieth Century Fruits. shipping facilities before,\" Burbank quipped to editors of his multi-volume biography, published in 1914.22 Garden and Forest recorded this rail-powered influence, announcing in 1895 that classic cultivars of domesticated European plum had arrived in New York on rail shipments from California, along with peaches, pears, and grapes. \"Forty-four car-loads of California fruits were sold here in five days of last week,\" the magazine reported.23 As the final achievement of the \"railroad era,\" Burbank--an alchemical breeder, dubbed a horticultural \"wizard\" in his own time24-- developed large thick-skinned plum hybrids that were easier to ship across country. Ironically, he used native species to impart that thicker skin, along with disease resistance, but otherwise, the flavor and appearance most strongly resembles the other parents: the Japanese plum (Prunus salicinia) and the apricot plum (Prunus simonii). 25 Burbank's hybrids--most famously `Santa Rosa'--still dominate the American plum industry and have been the parents of other successful and widespread cultivars.26 Another Era for the Native Plum Recent attempts to introduce native plums into the market have centered on beach plum (Prunus maritima), a species that hems the sandy coastline of New England, growing on shifting dunes, alongside American beachgrass (Ammophila breviligulata). The James R. Jewett Prize was established at the Arnold Arboretum for research on the species in 1940, and although the award waned after little more than a decade, it was reinstated in 1997 when researchers at Cornell University launched a concerted commercialization project.27 Richard Uva, who instigated the Cornell research under the direction of Professor Thomas Whitlow, now grows three acres of beach plums on his cut-flower nursery, Seaberry Farms, in Federalsburg, Maryland. He estimates that twenty-two acres of beach plum are currently under production between sixteen growers in Massachusetts, Maryland, New Jersey, and Long Island. This is not enough to meet the growing commercial demand, especially among distillers, brewers, and winemakers. Growers have also found a market among chefs, especially in tourist beach towns, where the plums are a stamp of local credibility for the menu. Jam and jelly productions remains a relatively small scale. For other plum species, production and research has yet to return, although interest in indigenous ingredients has swelled more generally in recent years. Part of this interest GREENHOOK GINSMITHS BIODIVERSITY HERITAGE LIBRARY 30 Arnoldia 75\/3 ? February 2018 One of the first of alcoholic products made from beach plum is a gin liqueur, manufactured by Greenhook Ginsmiths in Brooklyn. It is made in the manner of sloe gin--a product made with a European native plum (Prunus spinosa)--by steeping whole plums in gin for a matter of months. MICHAEL S. DOSMANN Native Plums 31 The author makes observations on a beach plum (Prunus maritima) submerged by sand dunes on Cape Cod in 2010. could be attributed to work by organizations like Slow Food, which, in 1996, launched a program known as the International Ark of Taste, designed to protect and preserve distinctive regional foods that are threatened with gastronomic extinction. Beach plums are listed among more than two hundred imperiled products in the United States, as are other oft-overlooked native foods like shagbark hickory nuts (Carya ovata), groundnut tubers (Apios americana), and tangy staghorn sumac fruits (Rhus thypina). Whether interest in native plums will be rekindled as part of this larger trend is yet to be seen. But during the historic boom of native plum cultivation, Abraham Dennis, an orchardist in Cedar Rapids, Iowa, became particularly inspired by the long history of plum cultivation among Native American communities in the region. He suggested there was almost a moral imperative to perpetuate the process. \"It is not alone our duty to rescue these fruits from their wild state and reawaken by culture these higher qualities given them by similar efforts by ancient horticulturists,\" Dennis said at a horticultural meeting in 1897. \"But,\" he continued, \"we must transmit them to future horticulturists more perfect fruits than we found them--new qualities added--worthy of the advanced and scientific age it is our privilege to live in.\"28 Now, well over a century later, Dennis's challenge resonates, enticing foragers and horticulturists back to the hedgerows and thickets to reclaim plums from the wild at last. Jonathan Damery is a former Curatorial Assistant and Curatorial Fellow at the Arnold Arboretum, and holds an MFA in creative writing from the University of Minnesota. His current book-length nonfiction project explores environmental history in the tallgrass prairie region, with grant support from the Minnesota State Arts Board in 2017. 32 Arnoldia 75\/3 ? February 2018 THE AMERICAN PLUMS Prunus americana by T. Davis Sydnor, The Ohio State University, Bugwood.org Plum taxonomy has long perplexed botanists, including Bailey. \"Native plums constitute probably the hardest [black] knot in American pomology,\" he wrote. \"The group is one of the most inextricably confused of any one of equal extent in our whole flora.\"29 More recently, Joseph Rohrer, writing about Prunus for the Flora of North America, described the \"particularly troublesome\" delineation of plum species. \"Surely,\" he wrote, \"as more molecular and genetic data are analyzed and, more importantly, correlated with morphological data, circumscription will be redrawn and the number of North American plum species further reduced.\"30 As it stands, Flora of North America recognizes thirteen species of native plums, outlined below. Prunus americana (American plum): The most widespread species, with a range stretching from New England to the Rocky Mountains. According to Hedrick, about 260 cultivars were derived from this species during the historic plum boom. Prunus angustifolia (Chickasaw plum): A distinctive southern species, ranging from Virginia to eastern New Mexico and south through Florida and other Gulf states. In 1911, Ulysses Hedrick counted about twenty horticultural selections of this species.31 Prunus geniculata (scrub plum): A federally endangered species found on sandy hills in central Florida. Fruits develop early compared to other species, from March to May. Prunus gracilis (Oklahoma plum): A small suckering species, no more than 1.5 m (4.9 ft) tall, which grows in dry, sandy locations. No significant horticultural varieties have been named, although Frank Waugh of the Vermont Agricultural Experimentation Station reported, in 1901, that the \"fruit is sometimes gathered and sold in local markets.\"32 Prunus angustifolia by Karan A. Rawlins, University of Georgia, Bugwood.org Prunus hortulana (hortulan or wild goose plum): Distribution for this species centers on Missouri and Illinois, with scattered pockets through southern Ohio and possible introductions in the Virginias. In older literature, this species was divided into two groups--wild goose and miner plums--which collectively resulted in about thirtysix named selections, according to Hedrick, although Kerr advertised more than fifty. Prunus maritima (beach plum): A denizen of the sandy coastline between Maine and Delaware. In 1911, Hedrick counted only two cultivars (`Bassett' and `Beta'). The num- Prunus nigra by R. W. Smith, Lady Bird Johnson Wildflower Center Native Plums 33 ber climbed by at least another dozen in the 1950s, through work funded by the Jewett Award, 33 and most recently, Rutgers released a cultivar named `Jersey Gem.' Prunus mexicana (Mexican or bigtree plum): The largest of native species, forming a tree up to 12 m (39 ft) tall. It ranges from northeastern Mexico to northern Illinois, east to Kentucky and Alabama. Prunus murrayana (Murray's plum): A suckering shrub known only from scattered populations near dry streams and canyon beds in southwestern Texas. Prunus nigra (Canada plum): A northern species that grows around the Great Lakes, ranging east to Massachusetts. It offered hardy characteristics for orchardists in Minnesota and Wisconsin, who, according to Hedrick, named about forty cultivars. Prunus subcordata by Terry Spivey, USDA Forest Service, Bugwood.org Prunus rivularis (creek or hog plum): A widespread species that has come to encompass a larger-statured taxon, Prunus munsoniana. The primary distribution is located between Texas and Missouri, with scattered populations to southern Ohio. At least sixty horticultural selections were made of this species by 1911, particularly among southern orchardists. Prunus subcordata (Sierra, Klamath, or Pacific plum): Found in California and Oregon, this is the only plum native west of the Rocky Mountains. Hedrick did not count the number of cultivars derived from this species, but he quoted Luther Burbank, who described certain selections with fruit as \"sweet as honey.\" Prunus texana by William R. Carr Prunus texana (peachbush or Texas wild peach): Long considered a peach rather than a true plum, this fuzzy-fruited species has a small range stretching from central Texas to the Gulf coast. DNA sequencing has revealed that subgeneric classification of Prunus is more complicated than the fiveparted system that formerly partitioned plums and peaches into separate subgenera. While other native peach-like species occur throughout the southwest, recent work places P. texana clearly within the plums.34 Prunus umbellata (hog, flatwoods, or Allegheny plum): A shrub or tree, growing to 6 m (19.7 ft) in height, this species ranges between North Carolina and eastern Texas. Traditionally a northern population in Michigan and the Allegheny Mountains was treated as a separate species, P. alleghaniensis, but the taxa have more recently been grouped. Prunus umbellata by James H. Miller and Ted Bodner, Southern Weed Science Society, Bugwood.org 34 Arnoldia 75\/3 ? February 2018 Endnotes, Native Plums 1Bradbury, J. 1817. Travels in the Interior of America in the Years 1809, 1810, and 1811. Liverpool: Smith and Galway, pp. 185?186. 2Whitson, J., R. John, and H. S. Williams (Eds.). 1914. Luther Burbank: His Methods and Discoveries and Their Practical Application, 1. London: Luther Burbank Press, pp. 275?77. 3Sargent, C. S. 1892. The Silva of North America: A Description of the Trees Which Grow Naturally in North America Exclusive of Mexico, 4. Cambridge: Riverside Press. p. 24 4Goff, E. S. 1891. The present status of native plum culture. Garden and Forest. 4 (193): 523?524. 5 Pe?a-Chocarro, L., et al. 2017. Roman and medieval crops in the Iberian Peninsula: A first overview of seeds and fruits from archaeological sites. Quaternary International: 1?18. 6See Goff, above. 7Bailey, L. H. 1892. The Cultivated Native Plums and Cherries. Ithaca: Cornell University Agricultural Experimentation Station. 8Ibid. 9Moerman, D. 1998. Native American Ethnobotany. Portland: Timber Press. 10Hedrick, U. 1911. The Plums of New York. Albany: L.B. Lyon Co. pp. 330?331. 11Lord, O. M. 1903. Minnesota City trial station. The Minnesota Horticulturist, 31(4): 130?131. 12Kerr, J. W. 1895. Price List of the Eastern Shore Nurseries: For Fall of 1895 and Spring of 1896. Denton, MD: Eastern Shore Nurseries. 13 Kerr, J. W. 1896. Annual Price-List and Catalog of the Eastern Shore Nurseries: Fall of 1896, Spring 1897. Denton, MD: Eastern Shore Nurseries. 14Kerr, J. W. 1909. Thirty-Seventh Year: Eastern Shore Nurseries and Fruit Farm. Denton, MD: Eastern Shore Nurseries. 15Wight, W.F. 1915. Native American Species of Prunus. Bulletin of the U.S. Department of Agriculture, No. 179. 16United States Census Office. 1900. Fruits, Nuts, and Forest Products. Agriculture, 5: 618. 17Ibid, 606. 181900. Plum, varieties--discussion. Annual Report of the State Horticultural Society of Missouri, 42: 222?223. 19See Kerr. 1896, above. 20See Hedrick, above. 21Lord, O. M. 1887. Native plums. Annual Report of the Minnesota State Horticultural Society, 15: 373?376. 22See Whitson and Williams, above. 231895. Notes. Garden and Forest, 393: 360. \"Notes.\" 24Janick, J. 2015. Luther Burbank: Plant breeding artist, horticulturist, and legend. HortScience, 50(2):153?156. 25See Whitson and Williams, above. 26Okie, W. R. and D. W. Ramming. 1999. Plum breeding worldwide. HortTechnology, 9(2): 162?176. 27Uva, R. H. 2003. Taming the Wild Beach Plum. Arnoldia, 62(4). 28Dennis, A. B. 1897. New native plums. Report of the Iowa State Horticultural Society for the Year 1896, 31: 79. 29See Bailey, above. 30Rohrer, J. 2014. Prunus. Flora of North America North of Mexico, 9. New York and Oxford: Flora of North America Association. 31See Hedrick, above. 32Waugh, F. A. 1901. Plums and Plum Culture. New York: Orange Judd Co. 33Garrick, L. 2012. Banking on beach plums. Gastronomica, 12(3): 21?30. 34Shaw, J. and R. Small. 2005. Chloroplast DNA phylogeny and phylogeography of the North American plums (Prunus subgenus Prunus section Prunocerasus, Rosaceae). American Journal of Botany 92:2011?2030. "},{"has_event_date":0,"type":"arnoldia","title":"Insights from a Sole Survivor: Quercus castaneifolia","article_sequence":9,"start_page":35,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25635","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15eaf6d.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Zale, Peter J.","article_content":"Insights from a Sole Survivor: Quercus castaneifolia Peter J. Zale O aks (Quercus spp.) have become a worldwide symbol of tree conservation. Of the approximately 450 described oak species, at least 175 are of conservation concern and many require further conservation assessments (Oldfield and Eastwood, 2007; Jerome et al., 2017). To aid in these efforts, public gardens around the world have invested significant energy to develop taxonomically and geographically exhaustive, genetically diverse oak collections with high conservation value. Aside from their importance as landscape and garden specimens, maintaining living collections of oaks in public gardens is particularly important because recalcitrant (desiccation intolerant) oak seeds cannot be stored long term in germplasm repositories, highlighting an increased need for ex situ conservation of individual trees and improvements in collection practices. Within the United States, the American Public Gardens Association (APGA) has recognized the efforts of 20 public gardens who have formed a Plant Collections Network Quercus multisite collection to tackle some of these efforts (APGA, 2017). Benchmarking studies have become an important tool to help these and other gardens prioritize their conservation and collecting efforts by revealing the diversity of oaks in their collections relative to those in cultivation elsewhere. Gaps, or missing species, reveal areas for future development, and the identification of species (or clones) uniquely or poorly represented in cultivation can indicate those in need of further preservation and distribution. However, the power of benchmarking is predicated on the quality of records and the verification of species to accurate identity. In 2015, verification of tree plantings on perimeter areas of Longwood Gardens revealed a mysterious oak bearing an incorrect label. An exercise in curatorial sleuthing led me on a chase to discover the species' identity to be Quercus castaneifolia, chestnut-leaf oak, and that the accession had a provenance linked to the wilds of northern Iran. In my further investigations, I learned more about its natural history and, eventually, saw it growing in its native habitat. Taxonomy and geography of chestnut-leaf oak Quercus castaneifolia was first described in 1831, and belongs to Section Cerris and Subsection Cerris, a placement within the genus that has remained stable since Camus' 1936 monograph. However, the taxonomy within Q. castaneifolia has been debated, with at least eight intraspecific botanical taxa (varieties, subspecies, and formae) described based on differences in leaf, acorn, trichome, foliar epidermis, and pollen morphology (Panahi et al., 2011). Molecular analysis using Amplified Fragment Length Polymorphisms (AFLP) indicated that the differences at the molecular level were not enough to distinguish among the previously proposed intraspecific taxa, except for Q. castaneifolia subsp. aitchisoniana, and that molecular variation in some cases was indicative of introgression from other species (Azadbakht et al., 2015). Presently, the botanical community considers Q. castaneifolia to be a single polymorphic or highly variable species (Rix and Kirkham, 2009; The Plant List, 2017). In addition to its distinct leaf characters, diagnostic morphological features of this species include its linear bud scales, and curiously elongated, but variably sized, ellipsoidal acorns--maturing in two years--that reach 2 to 4.5 cm (0.78 to 1.78 in) in length and have cupules or caps covered with prominent scales (see inside front cover). The branching habit of mature trees is variable, ranging from upright with a dominant central leader, to a spreading structure devoid of a strong central leader that LONGWOOD GARDENS 36 Arnoldia 75\/3 ? February 2018 Quercus castaneifolia is aptly named, with leaves [7 to 20 cm (2.8 to 7.9 in) long and 3 to 6 cm (1.2 to 2.4 in) wide] resembling those of the sweet or Spanish chestnut (Castanea sativa). This herbarium voucher was collected from the specimen growing at Longwood Gardens. Quercus castaneifolia 37 climate of Iran's Alborz Mountains (Panahi et al., 2011). In some of these areas, Quercus castaneifolia can form pure stands, although it becomes a member of Fagus orientalis (oriental beech)-dominated forest above 1200 m (3937 ft). It is a minor forest component in the eastern portion of its range, in areas dominated by Platycladus orientalis (Oriental arborvitae) (Menitsky, 2005). One study indicated that Q. castaneifolia comprises 6.5% of the total Hyrcanian Forest, belying the fact that it is among the most commonly encountered species in the region (Panahi et al., 2011). It has yet to be analyzed for Red Listing to determine its conservation status (Oldfield and Eastwood, 2017). The Specimen at Longwood Gardens The natural range of Quercus castaneifolia (shown in green) extends from Azerbaijan's Talysh Mountains along the southern border of the Caspian Sea to Gorgan, Golestan Province, Iran (map modified from Sales and Hedge, 1996). results in a distinctly rounded canopy. In the wild trees can reach heights of 45 m (148 ft), but 10 to 25 m (33 to 82 ft) tall and wide seems more typical, particularly in cultivated trees. Fall color develops late in the season in hues of clear yellow and brown. Chestnut-leaf oak is a member of the Hyrcanian Forest (from \"Hyrcania\", the Greek form of an old Persian word describing the region of Gorgan or Asterabad, Iran), a relict forest widespread during the Tertiary Period (65 to 15 million years ago) and now occurring only on the mountain ranges that surround the southern Caspian Sea. The forest is a well-defined glacial refugium rich in endemic species, including this oak (Milne and Abbott, 2002). Throughout its range, it grows from sea level to 2400 m (7875 ft) in elevation, and is reported to grow on the northern aspects of mountain slopes. (My personal observations, however, indicate a more general distribution, at least in Azerbaijan.) In the Talysh Mountains of southeastern Azerbaijan and northwestern Iran, it is a component of the Quercus-Buxus Forest that once dominated the Caspian Coastal Plain, and the Quercus-Carpinus Forest in the lowlands to 1200 m (3937 ft). The latter forest type transitions to the Quercus-Zelkova Forest in the drier In the 1950s Longwood Gardens became part of a large-scale project called the Michaux Quercetum. This project began as a partnership between the Morris Arboretum of the University of Pennsylvania and the Northeastern Forest Experiment Station of the USDA, and was financed in part by the Michaux Fund of the American Philosophical Society. The primary purpose of the study was to develop a large-scale provenance test of US native oak species with the goal of selecting genetically superior trees for breeding and forestry purposes. Another part of the project was defined loosely as \"preliminary tests of exotic oak species from all temperate oak-inhabited parts of the northern hemisphere\" (Schramm and Schreiner, 1954). Little information exists on the full extent of exotic species of oaks trialed in the various Querceta, but in 1968, several exotic oaks were planted adjacent to the original Michaux Quercetum here in Longwood Gardens. Among these were oak seedlings bearing USDA Plant Introduction (PI) numbers 228074 and 228075, both originally collected in 1955 as acorns by USDA Agricultural Explorer Howard Scott (H. S.) Gentry in northeastern Iran. The Plant Introduction Inventory (USDA, 1964), records Gentry's description for the collections, each listed as yet-to-be-identified Quercus sp. For PI 228074: \"Col. Nol. 15709. Twenty-five miles south of Kalow, Caspian slope of Alborz. Sept. 12, 1955. Elevation 6,000 feet. Second PETER ZALE 38 Arnoldia 75\/3 ? February 2018 In 2016, the specimen at Longwood Gardens (accession 1957-2444*A) was 15.5 m (50.9 ft) tall and 21.9 m (71.9 ft) wide, and had a DBH (diameter at breast height) of 112.5 cm (44.3 in). It has an ascending branching pattern and a distinctly rounded, spreading crown. growth from cut trees.\" And, for PI 228075: \"Col. No. 15799. Fifty-four miles east of Gorgan. Sept. 15, 1955. Spreading tree to 30 feet high.\" These were grown at the US Plant Introduction Garden (USPIG) in Glenn Dale, Maryland, and in September 1957 Longwood Gardens received three seedlings of PI 228074 and two of 228075, which were planted in the research nursery and later near the Michaux Quercetum. The Michaux Quercetum and adjacent oak plantings received little attention for the next five decades, providing a true test of a tree's ability to survive with relative neglect. During the summer of 2015, the plant records office was inventorying trees in this part of Longwood Gardens and `rediscovered' an unidentified oak tree. It was then brought to my attention and I began the detective work. An old planting map indicated the tree was Longwood Gardens accession 1957-2444*A, and the original pack- ing slip from the USPIG linked that accession to one of Gentry's collections: PI 228075 (accession 1957-2443 was assigned to PI 228074). It appears as if one seedling of each collection perished in the first few years. In 1971, an inventory of the Quercetum indicated that the two remaining seedlings of PI 228074 had perished due to sun scald, but no update was given for PI 228075. Using Gentry's original collection notes of the tree's nativity, I used the distinct shape of the tree's leaves, acorns, and linear bud scales to confirm the identify as Q. castaneifolia. Further research indicated that it is the only remaining tree in cultivation from Gentry's collection, and is thus unique among cultivated accessions of this species in public gardens worldwide. Because of its unique lineage, and the rarity of wild-collected Q. castaneifolia in the US (see below), I collected acorns in 2016. Oaks are TONY KIRKHAM Quercus castaneifolia 39 The Kew specimen (accession 1969-15985) is one of the largest trees at Kew, and is the largest chestnut-leaf oak in the British Isles. In 2016 it was approximately 35 m (115 ft) tall, 30 m (98 ft) wide, and had a DBH of 2.52 m (8.26 ft) at 1.5 m (4.9 ft). 40 Arnoldia 75\/3 ? February 2018 anemophilous (wind-pollinated) species that generally rely on fertilization from genetically different individuals of the same species for successful seed development. But there is some evidence that oaks have the ability for selfpollination on a limited basis (Yacine and Bouras, 1997). Of the 24 seeds collected, 12 germinated and the resultant seedlings so far appear to be true-to-type, despite extensive nearby plantings of US native and exotic species including Quercus macranthera (Caucasian oak), which is reported to hybridize with Q. castaneifolia. Despite promising initial results, verification to identity will have to wait until the seedlings mature. If anything, they can serve as understock to graft scions from the original tree, which would preserve the exact genetic lineage. Chestnut-leaf Oak in Cultivation The Botanic Gardens Conservation International (BGCI) PlantSearch website indicated that 68 public gardens worldwide (mostly in Europe) grow this species. For comparison, 148 gardens list the commonly grown Quercus cerris (Turkey oak), while 54 gardens list the less common Quercus libani (Lebanon oak). Perhaps the best-known specimen in cultivation is the tree growing behind the Water Lily house at Kew Gardens, London. Widely accepted as the first introduction of the species to cultivation in western Europe, the accession was received as seed in 1843 and was reputedly planted in 1846 by William Hooker himself (Rix and Kirkham, 2009). This imposing specimen is one of the most recognized trees in the collection at Kew and was one of few to survive the great storm of 1987 without damage. Records of this species in cultivation in the United States are few. Some of the largest are two 1938 trees (239-38*A and D) at the Arnold Arboretum of Harvard University, which came from the Mount Mashuk Forest Garden Experimental Station in Pyatigorsk, Russia (Northern Caucasus), which is outside of the species natural range. The larger of the two (239-38*D) stands 22.33 m (73.3 ft) tall, has a spread of 18.3 m (60 ft), and a DBH of 98.6 cm (38.8 in). A query of the most current inventory (2014) of the members of the Plant Collections Network multisite Quercus group indicated 19 living accessions of Quercus castaneifolia in nine gardens. Of these, 16 are of garden or nursery origin, two are from wild collections, and one is actually a hybrid: Q. castaneifolia ? Q. cerris. Interestingly, the two wild plants are 1994 accessions growing at the UC Davis Arboretum and originally came from Dr. Ahmad Mossadegh, Professor of Silviculture at the University of Tehran. He collected seeds from the Loveh Region, near Gorgan, Iran at an elevation of 830 m (2723 ft). This source locality, similar to that of the Longwood specimen, indicates that all of the known wild-sourced material in cultivation in the US comes from a similar place in the eastern extent of the species range. In addition to growing in the institutions mentioned above, chestnut-leaf oak is also at the Bartlett Tree Research Arboretum (Charlotte, NC), Denver Botanic Garden, Cornell Botanic Gardens (Ithaca, NY), and Morton Arboretum (Lisle, IL). Its ability to grow in such diverse places suggests a tolerance to extremes of heat and cold (USDA Hardiness Zones 5 to 9), as well as drought and a range of soil types. It is worth experimenting growing the tree in colder and drier regions where the palette of available landscape trees is limited. Quercus castaneifolia, recognized for its durability and ornamental qualities, has a number of selections. Aim?e Camus, in her 1936 Les Chenes: Monographie du genre Quercus, was the first to mention four cultivars (then formae) of Q. castaneifolia: `Asplenifolia', `Filicifolia', `Aureovariegata', and `Pyramidalis'. Although there were no descriptions provided, it is likely that the first two had fern-like, and the third had variegated leaves; the fourth likely had a narrow, yet not fastigiate, habit. No plants bearing these names have been found living in modern collections. Several other cultivars in Europe bear the place name where the original plant was selected. `Sopron', `Zorgvlied', and `Zuiderpark' originated in European parks or cities, and have not been widely propagated or distributed. Though listed as a selection of chestnut-leaf oak, `Algerensis' appears to be synonym for Quercus afares, the Algerian oak, a morphologically similar species from the coastal Atlas Mountains of Algeria and Tunisia. Quercus castaneifolia 41 PETER ZALE Perhaps the most common cultivar is `Green Spire' (note spelling, as numerous sources have it listed as `Greenspire'), selected and introduced by Hillier and Sons Nursery (Winchester, UK). It is described as \"a broadly columnar form of compact habit, raised in our nurseries about 1948. A vigorous, tall tree. Probably Q. castaneifolia ? Q. libani\" (Hillier Nurseries, 1991). This selection has become available in the US and its putative hybrid origin requires verification. The species is known to hybridize with others (particularly within Section Cerris), and although hybrids are poorly represented in cultivation, could prove valuable for creating widely adaptable, drought-tolerant trees suitable for managed landscapes, notably urban forests. Oikos Tree Crops (Kalamazoo, MI) offers Q. A large tree near Lankaran, Azerbaijan, had grey-brown bark with prominent, vertical ridges. castaneifolia ECOS Form, an open-pollinated strain sold as both seeds and seedlings from their original trees. According to their website description, the trees were originally obtained by growing acorns from gardens and arboreta, and may represent hybrids between Q. castaneifolia, Q. cerris, and Q. acutissima (sawtooth oak). They were reported to be the fastest growing oaks in their nursery, reaching nearly 2 m (6.6 ft) in height after two years from seed. Quercus castaneifolia in Azerbaijan From 9 to 22 September, 2017, the Plant Collections Collaborative (PCC, 14 US public gardens with similar interests in domestic and international plant exploration) organized and performed a plant collecting trip to the Republic of Azerbaijan. Participants included Phil Douglas (Chicago Botanic Garden), Matt Lobdell (Morton Arboretum), Vince Marrocco (Morris Arboretum of the University of Pennsylvania), and myself. Henrik Sj?man of the Gothenburg Botanical Garden (Sweden) joined us for the last week of the trip. Among the collection targets developed during trip planning was Quercus castaneifolia, no doubt inspired by my now familiarity with Longwood's single specimen. We were pleasantly surprised to find the species ubiquitous in the Lankaran Region of southern Azerbaijan, in the wild as well as a cultivated tree. It grew at sea level and was still abundant at 1500 m (4921 ft) in elevation, the highest point we reached between Lerik and Orand. We found it with other characteristic species of the Hyrcanian Forest: Acer velutinum (velvet maple), Buxus sempervirens (common boxwood), Carpinus betulus (European hornbeam), Parrotia persica (Persian ironwood), and Zelkova carpinifolia (Caucasian zelkova). Trunks of old, open-grown trees were often massive and supported a distinctive branching structure with a tall, straight central leader and irregular branches that resulted in a loosely rounded crown. Unfortunately, none of the trees bore mature acorns, and our guide, Dr. Ha?iaga Sofarov, Deputy-Director of Hyrcan National Park in Lankaran, indicated that they would not be ready until late-October or early-November. This corresponds to the timing of seed maturation of the tree at Longwood, but differs from 42 Arnoldia 75\/3 ? February 2018 HENRIK SJ?MAN ernmost extent of its range will increase the presence of wild-origin Q. castaneifolia at public gardens across the US, Conclusion Curatorial mysteries, like the story of Gentry's unidentified oak, are found in public gardens throughout the world. When acknowledged and solved, they not only enrich the collections and institutions where they exist, but also the greater public garden, horticulture, and botanical communities. As is the case for Quercus castaneifolia at Longwood Gardens, unraveling this mystery helped us inform, revise, and add value to our plant collections, their data, and our broader collections development objectives. The project shed new light on an underrepresented and under-collected species worthy of greater attention, and I hope this work serves to open new avenues of germplasm preservation, acquisition, and interpretation. References APGA (American Public Gardens Association). 2007? 2017. The plant collections network Quercus multisite group. Available at: https:\/\/ publicgardens.org\/programs\/plant-collectionsnetwork\/collections-showcase\/quercusmultisite. (Accessed 30 November 2017). The acorns found on trees near Lankaran, Azerbaijan were unripe in mid-September of 2017, foiling the collectors' plans. Gentry's field collection date of September 15, 1955, suggesting that acorns may mature earlier in the southeastern part of the species range. Chestnut-leaf oak is perhaps the most widely cultivated tree in the Lankaran Region. We immediately encountered extensive plantings used as windbreaks amongst the vast agricultural expanses that dominate the Caspian Lowlands. Street trees were also common, which thrived despite compaction from surrounding sidewalks and streets, late-summer heat and drought, pollution from vehicle exhaust and general neglect. PCC members are again planning an autumn 2018 trip again to southern Azerbaijan to make collections of this and other important, under-represented species. Hopefully, our success in capturing the northwest- Azadbakht, M., I. Mehregan, M. Assadi and T. Nejadsattari. 2015. Phylogenetic Relationships in Quercus castaneifolia C.A. Mey. in Hyrcanian forests of Iran based on AFLP markers. Journal of UMP Social Sciences and Technology Management 3:237?246. Bean, W.J. 1976. Trees and shrubs Hardy in the British Isles, Volume III. 8th revised edition. Butler and Tanner Limited, London, U.K. Camus, A. 1936. Les Chenes: Monographie du genre Quercus. Paul Lechevalier and Fils, Paris, France. Hillier Nurseries. 1991. Hillier's Manual of Trees & Shrubs. David and Charles, London. Jerome, D., E. Beckman, L. Kenny, C.S. Kua, and M. Westwood. 2017. The Red List of U.S. Oaks. The Morton Arboretum, Lisle, IL. Kr?ssman, G. 1978. Manual of Cultivated Broad-leaf Trees and Shrubs. Volume III, PRU-Z. Timber Press, Portland, Oregon, U.S. Lancaster, R. 1974. Paradise found. Journal of the Royal Horticultural Society 99:103?109. PETER ZALE Quercus castaneifolia 43 A restaurant nestled in a shelterbelt of chestnut-leaf oak north of Lankaran, Azerbaijan. Like cultivated trees observed elsewhere, the trunks were painted white, supposedly as a preventative measure against trunk damaging insects. Menitsky, Y.L. 2005. Oaks of Asia. Science Publishers, Enfield, New Hampshire, U.S. Milne, R.I. and R.J. Abbott. 2002. The origin and evolution of Tertiary relict flora. Advances in Botanical Research 38:281?314. DOI: 10.1016\/S00652296(02)38033-9 Oldfield, S. and A. Eastwood. The Red List of Oaks. Flora and Fauna International, Cambridge, UK. Panahi, P., Z. Jamzad, M. R. Pourmajidian, A. Fallah, and M. Pourhashemi. 2011. A revision of chestnutleaved oak (Quercus castaneifolia C. A. Mey.; Fagaceae) in Hyrcanian Forests of Iran. Caspian Journal of Environmental Sciences 9:145?158. Rix, M. and T. Kirkham. 2009. Quercus castaneifolia. Curtis's Botanical Magazine 26:54?63. Sales, F., and I.C. Hedge. 1996. Biogeographical aspects of selected SW Asiatic woody taxa. Annalen des Naturhistorischen Museums in Wien. 98B Supplement:149?161. Schramm, J.R. and E.J. Schreiner. 1954. The Michaux Quercetum. Morris Arboretum Bulletin 5:54?57. Snyers, C. 2014. A natural hybrid? Available at: http:\/\/ www.internationaloaksociety.org\/ natural-hybrid\/ (Accessed: 30 November 2017) The Plant List. 2017. Version 1.1. Published on the I n t e r n e t ; h t t p : \/ \/ w w w. t h e p l a n t l i s t . o r g \/ (Accessed: 28 November 2017). Trehane, P. 2007?2015. The Oak Names Checklist. Available at: http:\/\/oaknames.org\/search\/ goodnames.asp. (Accessed: 30 November 2016). USDA (U.S. Dept. of Agriculture). 1964. Plant Inventory No. 163. Plant Industry Station, Beltsville, Maryland, U.S. Yacine, A. and F. Bouras. 1997. Self- and cross-pollination effects on pollen tube growth and seed set in holm oak Quercus ilex L. (Fagaceae). Annals of Forestry Science 54:447?462. Peter Zale, Ph.D., is the Associate Director for Conservation, Plant Breeding, and Collections at Longwood Gardens in Kennett Square, PA. "},{"has_event_date":0,"type":"arnoldia","title":"A Winter Beauty: Viburnum opulus","article_sequence":10,"start_page":44,"end_page":45,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25631","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d15ea726.jpg","volume":75,"issue_number":3,"year":2018,"series":null,"season":null,"authors":"Keegan, Brendan","article_content":"A Winter Beauty: Viburnum opulus Brendan Keegan W hen I think back on my experience as a Peace Corps volunteer in Ukraine, my memories are often drawn to the cold, snowy days of January. In the silence of winter afternoons, the plants in the small town of Terebovlya seemed especially distinct. I often walked through haunting groves of silver birch (Betula pendula), white bark against the white snow of fields beyond. Near the school where I worked, two magnificent bigleaf lindens (Tilia platyphyllos) towered like giants among young European hornbeam (Carpinus betulus). However, nothing stood out against the winter gray so much as the bright red fruits of Viburnum opulus, a plant that Ukrainians have long praised in song, poem, and prose as a visceral symbol of beauty and identity. Native to Europe, Asia, and North Africa, Viburnum opulus is a multi-stemmed deciduous shrub. It has a rounded growth habit and can grow up to 5 meters (16 feet) tall. Although it is known as kalyna in Ukrainian, Western Europeans often call it the guelder rose, so-named for a region of the Netherlands where the popular \"snowball tree\" cultivar supposedly originated. In North America, Viburnum opulus is called European cranberrybush, because of its tart, cranberry-like red fruits, despite the fact that it is in the moschatel family (Adoxaceae) and unrelated to the true cranberry (Vaccinium macrocarpon), a member of the rhododendron family (Ericaceae). (European cranberrybush is V. opulus var. opulus; the similar looking American cranberrybush, previously known as V. trilobum, is now known as V. opulus var. americanum.) During the summer months, Viburnum opulus bears three-lobed, dark green leaves, 5 to 10 centimeters (2 to 4 inches) long and wide. These palmate leaves, which resemble those of some maples, have deeply impressed venation, wrinkled surfaces, and soft undersides. In the autumn, the foliage often turns beautiful hues of red and purple. Red is also the color of the fully ripened berry-like drupes (fleshy, single-seeded fruits), which mature in late fall and can remain on the plant until the following spring. The vibrant fruits hang in dense clusters and, although they are primarily consumed by birds, they are also edible to humans. Tart and bitter until softened by frost, they are nonetheless believed to have medicinal properties, and Ukrainians consume small quantities raw, baked, or in tea to help treat various illnesses. It is also common to see fruit clusters adorning entryways, as well as on traditional Ukrainian embroidered clothing, as symbols of health and fertility. In North America, the beautiful white lacecap inflorescences of Viburnum opulus are often considered the plant's defining aesthetic characteristic. Each cyme is composed of a single ring of large, white petaled, sterile florets on the outside and bunches of smaller fertile florets on the inside. Some cultivars, such as the popular V. opulus `Roseum' produce inflorescences composed of entirely sterile flowers that look like snowballs or pom-poms, leading to the nickname \"snowball tree.\" Another popular cultivar, Viburnum opulus `Xanthocarpum' has the typical flat white corymbs, but produces bright golden yellow fruits instead of red. Earlier this year, I stopped by three Viburnum opulus accessions (352-78*A, C, and E, collected in the wild from the northeast of Denmark) in our Viburnum Collection. Humble looking among their neighbors, it was interesting to reflect that this plant is ubiquitous in small Ukrainian towns and villages where it is proudly planted next to homes. I had the chance to reflect on this again later in October when my wife and I were visiting friends in Ukraine. While there, one of her former colleagues asked where I worked, and I stumbled over describing the Arnold Arboretum and its mission of plant research and conservation. However, when he followed up my explanation by asking whether or not we grew kalyna, I was proud to say yes. Brendan Keegan is a Gardener at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23460","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14eb328.jpg","title":"2018-75-3","volume":75,"issue_number":3,"year":2018,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Food, Poison, and Espionage: Mycorrhizal Networks in Action","article_sequence":1,"start_page":2,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25627","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14e856b.jpg","volume":75,"issue_number":2,"year":2017,"series":null,"season":null,"authors":"Yih, David","article_content":"Food, Poison, and Espionage: Mycorrhizal Networks in Action David Yih C ROB ROUTLEDGE, SAULT COLLEGE, BUGWOOD.ORG an trees nurse their young? Do plants send out signals underground to warn each other of the arrival of ravenous insects? Can they go on the attack themselves and cripple competing plants with noxious chemicals they deliver through fungal connections? Lately, researchers investigating subterranean fungal networks have come up with surprising answers to questions like these. Their intriguing findings also have game-changing implications for ecology and conservation, forestry and agriculture--even evolutionary theory. Some 90% of terrestrial plant species around the world engage in symbioses called mycorrhizae--from Greek mykos (fungus) and rhiza (root). Mycorrhizal plants come from all corners of the plant kingdom and include trees, forbs, grasses, ferns, clubmosses, and liverworts. Their symbiotic partners (symbionts) are fungi whose threadlike hyphae radiate out into the soil, bringing water and nutrients--including phosphorus, nitrogen, zinc, and copper--back to the plant's roots in exchange for a share of the carbohydrates plants produce through photosynthesis. Though mycorrhizal symbioses range along a continuum from parasitic (on the part of the plant) to mutualistic types, most are mutually beneficial. By themselves, plants can only access nutrients in the immediate vicinity of their feeder roots, and soon exhaust the supply. By associating with fungi, they conserve resources that would have been spent on growing ever larger root systems. In fact, as Smith and Read state categorically in their compendium of all things mycorrhizal, \"Mycorrhizas, not roots, are the chief organs of nutrient uptake by land plants\" (Smith and Read 2008). Subway Lines The native North American orchid, hooded coralroot (Corallorhiza striata), is a mycoheterotroph that relies on mycorrhizal networks for its survival. When compatible mycorrhizal fungi and plants recognize each other and create an interface in the plants' roots for the exchange of nutrients, they can form a variety of structures in and around the roots, depending on the species involved. Broadly speaking, there are two main types: arbuscular mycorrhizae and ectomycorrhizae. Arbuscular mycorrhizae get their name from the classic shape that the fungi take inside root cells, a profusely branching form called MARK BRUNDRETT an arbuscule. As its name suggests, an arbuscule has a miniscule tree- or shrub-like shape. The creation of an arbuscular mycorrhiza begins when chemicals exuded by a plant's roots stimulate a nearby arbuscular mycorrhizal fungus to branch and grow, allowing it to quickly find the roots. Once in contact, the fungus adheres to the root surface and, within a few days, penetrates the root and begins the formation of a mycorrhiza. Inside the root, different arbuscular mycorrhizal structures may develop, This tree-shaped arbuscule within a plant cell is part of an arbuscular mycorrhizal depending on the particular fungus in the genus Glomus. species involved. In 1905, the botanist Ernest-Isidore Gallaud named arbuscular mycorrhizal structures after plant genera he found them in. Arum-type mycorrhizae resemble maps of bus or subway lines: hyphae grow into the space between rows of cells, extending alongside them, like avenues running past city blocks, and making \"stops\" along the way to enter cells and form arbuscules. Though it penetrates a root cell's wall, the fungus remains in what amounts to an antechamber; it never passes through the cell's plasma membrane. Instead, Both coils and small arbuscules can be seen within these root cells of yellow trout this membrane envelops the lily (Erythronium americanum). invading hypha and all of its branches, maximizing the area of mutual conParis type, the fungus forms coils that look like tact. At this interface, plant and fungus estabchaotic loops of strewn intestines. An occalish a sort of marketplace where each partner sional small arbuscule may branch off from deposits nutrients and trades them for nutrients a coil, but the Paris-type mycorrhiza lacks a deposited by the other. straight \"subway line\" traveling alongside the Gallaud named the other main form of arbuscells. Instead, a coiling hypha exits from one cular mycorrhiza the Paris type, after a Eurcell only to enter the adjacent one, where it asian plant genus (a relative of Trillium). In the forms another mass of coils before moving on MARK BRUNDRETT Mycorrhizal Networks 3 MARK BRUNDRETT to the next cell--definitely not the express train! Arbuscular mycorrhizae have been around for a long time. Researchers have found arbuscules in fossils of Aglaophyton, an extinct genus of pre-vascular plant, dating from around 410 million years ago, in the Devonian Period. In fact, most scientists agree that the first plants to colonize the land were symbiotic organisms containing structures very similar to arbuscular mycorrhizae (Smith and Read 2008). Ectomycorrhizal roots of a container-grown conifer. Given their lengthy tenure on the planet, it's not surprising that arbuscular mycorrhizae occur in nearly all species of herbaceous plants and in most trees and shrubs. But the other main mycorrhizal type--the ectomycorrhiza (going back a mere 50 million years)--is also extremely important. While only about 3% of seed plants are ectomycorrhizal, they occupy large expanses of the earth's terrestrial surface. Almost all are woody plants, and they include forest trees that are the world's main sources of timber, such as pines. Starting alphabetically, Abies, Acer, The Hartig net of an ectomycorrhizal fungus extends among poplar (Populus) root cells. Alnus, Betula, Carpinus, and grows between the outer layers of the root's Corylus are a few of the eastern North American genera containing at least one ectomycorcells. Another difference involves the relative rhizal species. numbers of associated fungus species. Despite Ectomycorrhizae differ from arbuscular the vast numbers of arbuscular mycorrhizal mycorrhizae in several other ways. Unlike plant species, their fungal symbionts consist arbuscular mycorrhizal fungi, ectomycorrhizal of only about 150 species, all in the division fungi mostly do not penetrate root cells, hence Glomeromycota. Inversely, a more diverse their designation as \"ecto-\" meaning \"outer\" group of about 5,000 to 6,000 fungus species or \"external.\" The hallmarks of an ectomyform ectomycorrhizal associations. And while corrhiza are the fungal sheath, which encloses all arbuscular mycorrhizal fungi are microscopic and subterranean, many ectomycorthe root tip in a dense mass of hyphae, and the rhizal fungi develop large fruiting bodies that Hartig net, a labyrinthine hyphal network that ROBERT L. ANDERSON, USDA FOREST SERVICE, BUGWOOD.ORG 4 Arnoldia 75\/2 ? November 2017 Mycorrhizal Networks 5 occur either above or below ground. Those that appear above ground include many common woodland mushrooms, while the most notable of the underground-fruiting ectomycorrhizal fungi belong to the genus Tuber, best known for its fruiting bodies, truffles. Sharing Food We can visualize a mycorrhiza as a simple oneto-one relationship between an individual plant and an individual fungus. But in nature the picture is more complex. As the threadlike hypha of a mycorrhizal fungus extends outward from a plant's roots, it frequently encounters the roots of other plants of the same or different species. It may form mycorrhizae with these new partners, while still maintaining its connection with the first plant. As it proliferates in new directions, the hypha branches and fuses repeatedly, weaving a fungal net through the surrounding soil. Meanwhile, additional fungi of the same or different species may approach the first plant. If they're compatible, the plant is apt to form mycorrhizae with them, too. Soon a diverse association appears, composed of various fungi and various plant species, big and small, all connected into a sizeable mycorrhizal network that may span hundreds of hectares of forest (Gorzelak et al. 2015). The promiscuous nature of these associations of multiple plant and fungus species has prompted scientists to give their papers playful titles like: \"Changing partners in the dark,\" \"Mycorrhizal networks: des liaisons dangereuses,\" and \"Architecture of the wood-wide web.\" Mycorrhizal networks are highly efficient at procuring essential plant nutrients from the soil while the plant partner provides the carbon that fungi require. But the carbon doesn't stop there. It's long been known that certain nonphotosynthetic, parasitic plants, called mycoheterotrophs, depend on carbon shuttled from photosynthesizing plants via mycorrhizal fungi (see next page). The seeds of most mycoheterotrophs are tiny \"dust seeds,\" consisting of only a few cells and little or no endosperm to supply the germinating plant with food. Thus, these species depend upon mycorrhizal fungi for their survival. The orchids, perhaps the largest family in the plant kingdom, depend entirely on carbon received via mycorrhizal fungi for successful seed germination and early development. Experiments have shown that certain green orchids can convey carbon back to their associated fungi once they reach maturity. Thus, they partake in mutualisms that are offset in time, like borrowers repaying a loan. But \"full mycoheterotrophs\" (including some orchids) depend, throughout their lives, on carbon received through mycorrhizal networks, apparently without benefit to the fungus. In recent years investigators have discovered that mycorrhizal networks can distribute resources in much more flexible ways than previously thought, sending them in the direction of greatest need in response to changing conditions, in a seasonal tide-like flux. Researchers at Laval University in Quebec found evidence that carbon moved via mycorrhizal networks from yellow trout lilies (Erythronium americanum) to young sugar maples (Acer saccharum) as the maples' leaves unfurled in spring, and then back to the trout lilies in the fall during rapid trout lily root growth (Lerat et al. 2002). The direction of carbon flow can reverse even more frequently. University of British Columbia researchers reported that the flow of carbon changed direction not once but twice in the course of a growing season. In the spring, carbon traveled from Douglas-fir (Pseudotsuga menziesii) to paper birch (Betula papyrifera) as its buds resumed growth. In the summer, carbon flowed from heavily photosynthesizing paper birch into stressed Douglas-fir in the understory. And in the fall, it flowed from stillphotosynthesizing Douglas-fir into paper birch as it shed its leaves (Simard et al. 2012). Other resources besides carbon can change direction too. Though water typically flows from mycorrhizal fungi into the roots of their plant symbionts, under extreme conditions it can go the other way. In a greenhouse experiment, investigators using dye tracers found that when soil became extremely dry, oaks that were able to access water through their deep taproots transferred water to their mycorrhizal fungi, thus keeping them alive (Querejeta et al. 2003). Plants can even defend their fungal partners from fungivores. A recent study found evidence that when springtails (tiny insect-like Mycoheterotrophs and the Birth of Mycorrhizology The first impetus for the investigations that culminated in the discovery of mycorrhizae in the nineteenth century came from a puzzling group of plants that grew in the gloom of the forest floor. The ghostly apparitions lacked chlorophyll and had only vestigial leaves. Instead of spreading deeply into the soil, their roots tended to be tangles of truncated \"coralloid\" stubs. How did such plants thrive in the darkness? In 1841, when a British entomologist and botanist named Edward Newman decided to publish a new monthly called The Phytologist, he had no idea that a controversy would arise among its pages that would mark the beginning of an entirely new field of inquiry. At the center of the debate was a peculiar plant that Linnaeus had named Monotropa hypopitys (now Hypopitys monotropa, yellow pinesap). With a wide distribution encompassing much of temperate Eurasia and North America, pinesap was one of the most common achlorophyllous plants in Britain, and its odd lack of green color cried out for an explanation. As early as 1821, William Jackson Hooker had posed the question of whether the plant was a parasite. Botanists had long been aware of the existence of parasitic plants, like those in the genus Orobanche, that attached themselves to other plants' roots. But the nature of pinesap's attachment, if any, to other plants' roots was unclear. In 1840, the Austrian botanist Franz Unger had looked at the closely intertwined roots of pinesap and Norway spruce (Picea abies) and concluded that it was not a parasite. A year later, the editor of The Phytologist proposed to its readers that they take up the question. The first response, from Edwin Lees, appeared in the December 1841 issue: the plant was a parasite on beech roots. Two issues later, another contributor came to the opposite conclusion: the plant was not a parasite. Soon more readers joined the fray, and the controversy raged on for months. The one thing the respondents could agree on was that they'd all seen an annoying profusion of mysterious fibers that impeded their attempts to observe any connection between tree roots and pinesap roots. The status of pinesap remained an open question until 1960, when Swedish investigator Erik Bj?rkman used the carbon-14 isotope to trace the movement of carbon into pinesap from nearby trees. Lees had actually hit the mark when he'd written that the strange fibers looked fungal and appeared to be \"imbibing nutriment from the rootlets of beech to which they are closely applied, and conveying it to the succulent radicles of the Monotropa, with which they are also connected.\" Pinesap turned out to be a previously unknown type of parasite that acquires carbohydrates from green plants by connecting to mycorrhizal networks. Scientists have dubbed such plants mycoheterotrophs and speculate that they evolved from photosynthetic mycorrhizal plants adapting to the darkness of the forest floor. Having no requirement for sunlight, most full mycoheterotrophs are subterranean for much of their lives. Some even flower and set seed underground! THOMAS G. BARNES, UNIVERSITY OF KENTUCKY hexapods) browse on mycorrhizal fungi, plants can help by sending protective chemicals into the hyphae (Duhamel et al. 2013). What about the extraordinary idea that plants might be subsidizing their progeny-- essentially nursing them-- using mycorrhizal networks? Though there is no clear evidence that plants can detect their kin through mycorrhizal networks and shuttle nutrients preferentially to offspring, there are hints in that direction. Ferns reproduce in a life cycle that passes through two distinct generations. Spores from the familiar, leafy sporophyte generation ger minate and grow into the tiny, rarely seen gametophyte generation, which, through sexual reproduction, gives rise to the next generation of sporophytes. Researchers working with two species of the fern genus Botrychium found that strains of Glomus (a genus of arbuscular mycorrhizal fungi) maintained mycorrhizae with individuals of both generation types simultaneously, demonstrating the potential for sporophytes to subsidize the Research showed that mycorrhizal networks reversed the movement of carbon between yellow trout lilies (Erythronium americanum) and sugar maples (Acer achlorophyllous gametophytes saccharum) during the year. (Winther and Friedman 2007). about? Like many animal species, plants have Since then, a number of studies have shown that tree seedlings do indeed benefit from a language of danger. In the early 1980s, David resources received from mature trees of the Rhoades, a zoologist interested in the interactions between insect herbivores and plants, same species via mycorrhizal networks, though proposed a novel idea. In the course of his not necessarily to a greater degree than other research with Salix sitchensis, he had noticed plants in the network. that defensive changes in the leaf chemistry of Notes from the Underground willows being chewed on by tent caterpillars The hustle and bustle of mycorrhizal netalso showed up in the leaves of nearby plants, works becomes even more intriguing as we even though they had not yet been attacked. look beyond resource sharing to the remarkHe speculated that the neighboring plants must able communication functions of mycorrhihave detected airborne molecules emanating zal networks. What do plants need to talk from either the attacked plants or the tent cat- JOSEPH O'BRIEN, USDA FOREST SERVICE, BUGWOOD.ORG Mycorrhizal Networks 7 8 Arnoldia 75\/2 ? November 2017 erpillars, prompting them to deploy protective chemicals preemptively (Rhoades 1983). Subsequent research confirmed Rhoades' suspicion-- plants being attacked by herbivores can release volatile organic compounds into the air that induce defensive responses in nearby plants. And recent experiments have shown that such \"stress signals\" can also be transmitted through mycorrhizal networks. Researchers at South China Agricultural University inoculated tomato plants with the fungal pathogen Alternaria solani, the cause of early blight disease in tomatoes and potatoes, and became the first to demonstrate that mycorrhizal networks can act as plant-to-plant communication conduits. They found that the uninfected tomato plants (stress-signal receivers) in the mycorrhizal network showed an increase in disease resistance and putative defense-related enzyme activity. They also found that the receiver plants had activated several defense genes. These changes in the receiver plants began within 18 hours of inoculating the donor plants (Song et al. 2010). In 2013, a group of scientists working in the United Kingdom decided to follow up on the fungal pathogen study and see what would happen with insect herbivores. They produced the first experimental evidence that signal molecules from plants infested with aphids travel through mycorrhizal networks to uninfested neighboring plants. Within 24 hours of the arrival of pea aphids (Acyrthosiphon pisum) on broad bean plants (Vicia faba), signals traveling through mycorrhizal networks caused uninfested broad beans to give off volatile compounds. Not only did these compounds repel the aphids, they actually attracted the aphids' natural enemy, the parasitoid wasp Aphidius ervi (Babikova et al. 2013). This interaction apparently benefits all three parties to the network. Being quickly alerted to the threat allows the uninfested beans to deploy their protective volatiles preemptively, thus evading aphid attack. The fungi thereby avoid a potentially catastrophic reduction in the plants' capacity to supply them with carbon. And even the infested beans may benefit: some investigators suggest that stress-signal transmission ensures that signal-donor plants will become engulfed in a large plume of protective volatiles created collectively by the surrounding plants in the network (Barto et al. 2012). In some cases, attacks on plants can simultaneously stimulate both stress signals and nutrient transfers. A recent collaboration between Chinese and Canadian researchers investigated the flow of carbon and stress signals in a mycorrhizal network involving a four-month-old interior Douglas-fir (Pseudotsuga menziesii var. glauca), a ponderosa pine (Pinus ponderosae), and the ectomycorrhizal fungus, Wilcoxina rehmii. They found that manual defoliation of the young Douglas-fir resulted in a transfer of both defense signals and carbon via mycorrhizal network to the ponderosa pine (Song et al. 2015). Some mycorrhizologists ascribe this result to the fungus throwing in its lot with the healthy pine rather than throwing good money after bad by propping up the struggling Douglasfir. Postulating that the transfers were initiated by the fungus, they write: \"Here, the networking fungus may have acted to protect its net carbon source, by allocating carbon and signals to the healthy, more reliable ponderosa pine\" (Gorzelak et al. 2015). Weapons of Plant Destruction Besides tranferring resources and signals, mycorrhizal networks can extend the reach of the allelochemicals that certain plants produce--toxic substances that inhibit the development of nearby competitors. Thus, \"mycorrhizal networks can serve as couriers for biochemical warfare\" (Gorzelak et al. 2015). A study of the effect of mycorrhizal networks in the transport of the allelochemical juglone, which is exuded by the roots of Juglans species (walnuts) and negatively affects the growth of many plants including rhododendrons, tomatoes, and apples, unequivocally implicated mycorrhizal networks in the dispersal of juglone into the soil (Achatz et al. 2014). In at least one case, instead of helping to spread noxious allelochemicals, mycorrhizal fungi themselves become the victims. Garlic mustard (Alliaria petiolata), a European plant well known as an invasive in eastern North America, is a non-mycorrhizal plant that produces fungicidal allelochemicals. Researchers found that garlic mustard drastically reduced the abil- JOSEPH BERGER, BUGWOOD.ORG Mycorrhizal Networks 9 the West, where Idaho fescue (Festuca idahoensis) is a common native grass. University of Montana researchers estimated that as much as 15% of the above-ground carbon in spotted knapweed plants came from nearby fescue by way of mycorrhizal fungi (Carey et al. 2004). Thus, invasives may exploit mycorrhizal networks to thrive at the expense of neighboring native plants. Networking for the Future ADOLF AND OLUNA CESKA Mycorrhizal networks can transmit warning signals to surrounding plants when pests such as these pea aphids attack. The fruiting bodies of the ectomycorrhizal fungus Wilcoxina rehmii. ity of North American arbuscular mycorrhizal fungal spores to germinate and form mycorrhizae. As a result, American mycorrhizal plants had reduced seed-germination and increased mortality, while non-mycorrhizal plants were unaffected. European arbuscular mycorrhizal fungi and plants were also relatively unaffected, presumably due to their long evolutionary exposure to garlic mustard's allelochemicals (Callaway et al. 2008). Another peculiar relationship between invasives and mycorrhizal networks involves spotted knapweed (Centaurea stoebe, formerly C. maculosa), which is invasive in many areas and covers over seven million acres in the United States. It's of particular concern in Understanding mycorrhizal networks is evidently important for effective conservation of many species. This is particularly true of mycoheterotrophs, which cannot survive apart from mycorrhizal networks. According to Martin Bidartondo of the Royal Botanic Gardens at Kew, \"myco-heterotrophic plants are excellent indicators of undisturbed forests and forests with old-growth characteristics\" (Bidartondo 2005). It follows that mycoheterotrophs are among the species at greatest risk of extirpation from the clearcutting of forest lands (Moola and Vasseur 2004). Mycoheterotrophs are extremely hostspecific, so their conservation must involve both their particular fungal host species and the green plants that supply carbohydrates as essential habitat components. With the increased resistance to diseases and pests and the better access to water and nutrients that mycorrhizal networks offer, there is increasing recognition of the potential for a new \"Green Revolution\" based on using mycorrhizae in crop fields and forests. Much of the world's agriculture depends upon fertilizer derived from mined rock phosphate, a non-renewable resource that is steadily dwindling. Phosphorus is a crucial plant nutrient that mycorrhizal fungi are particularly good at locating in ordinary soil and funneling back to their plant symbionts. We can lessen our dependence on rock phosphate by finding ways to work with mycorrhizae. Proposed techniques include sowing fallow fields with appropriate mycorrhizal plants to maintain the level of fungal inoculum in the soil between crop rotations, using tilling patterns that minimize disturbance of mycorrhizal fungi, and avoiding the indiscriminate use of fungicides in the soil. Many tree nurseries are finding that inoculat- Garlic mustard (Alliaria petiolata) not only crowds out woodland natives but also exudes allelochemicals that negatively affect arbuscular mycorrhizal fungi. EDWARD L. BARNARD, FL DEPT OF AGRICULTURE AND CONSUMER SERVICES, BUGWOOD.ORG ing tree seedlings with appropriate mycorrhizal fungi increases survival both in the nursery and after planting out. In perusing the reports mentioned in this article, I was struck by the various ways investigators conceptualized what they saw happening in mycorrhizal networks. There are large gaps in what is understood about how mycorrhizae operate, and scientists must often use human metaphors as stand-ins to bridge the gaps. One implicit question that kept surfacing was: Who were the doers of the actions taking place in mycorrhizal networks, and what were their \"motives\"? Were plants \"nursing\" their progeny to keep their species going, or were fungi redistributing resources to the young plants with an eye to their own future wellbeing? Were Douglas-firs helping paper birches so as to later receive reciprocal benefits in their hour of need, or were fungi orchestrating the flux of resources, minimizing their risk by diversifying across multiple partner species? Were stress-signal donors \"warning\" receiver plants, or were the receivers \"eavesdropping\" on donors, on the alert for potential trouble? Or were mycorrhizal fungi acting like savvy farmers, apportioning fertilizer and coordinating pest management to maximize long-term yield? Perhaps the answer is \"all of the above,\" because ultimately all the organisms involved tend to strengthen and perpetuate their mutually beneficial networks. Indeed, when all the participants' roles are considered, the network as a whole emerges as a kind of higherorder organism in its own right, fitter than the sum of its parts, a well-ordered social entity capable of surviving the death of any of its individual members. Some scientists argue that the groupings of species involved in mycorrhizal networks are examples of natural selection at the level of the group (Gorzelak et al. 2015). For others, the interesting question is: which is the true driver of evolution--competition or cooperation? The ground-breaking evolutionary theorist Lynn Margulis passionately insisted on the predominant role of symbiosis in evolution. And for evolutionary biologist and author Frank Ryan, the discovery of mycorrhizae was a missed opportunity. He wrote, \"The NANCY ROSE 10 Arnoldia 75\/2 ? November 2017 The ectomycorrhizal fungus Pisolithus tinctorius readily makes associations with a number of woody plants and is used in plant nurseries to improve seedling growth. Seen here, a young fruiting body (left) and associated mycorrhizal roots on slash pine (Pinus elliottii) seedlings. intimate cooperation between wholly different life forms--plants and fungi--is not only an amazing biological phenomenon but also a vitally important factor in the diversity of plant life on earth. It should have been of enormous interest to evolutionary theorists, Mycorrhizal Networks 11 but ... at the end of the nineteenth century, as the fundamental principles of biology were being hammered into place in laboratories around the world, Darwinian evolution took center stage. And as Darwinism, with its emphasis on competitive struggle, thrived, [mutualistic] symbiosis, its cooperative alter ego, languished in the shadows, derided or dismissed as a novelty\" (Ryan 2002). Perhaps its time is still to come. In the meantime, plants and their mycorrhizal networks offer a fascinating and fruitful field of inquiry on many different levels. References Achatz, M., E. K. Morris, F. M?ller, M. Hilker, and M. C. Rillig. 2014. Soil hyphamediated movement of allelochemicals: arbuscular mycorrhizae extend the bioactive zone of juglone. Functional Ecology 28: 1020?1029. Babikova, Z., L. Gilbert, T. J. A. Bruce, M. Birkett, J. C. Caulfield, C. Woodcock, J. A. Pickett, D. Johnson, and N. van Dam. 2013. Underground signals carried through common mycelial networks warn neighbouring plants of aphid attack. Ecology Letters 16: 835?843. Barto, E. K., J. D. Weidenhamer, D. Cipollini, and M. C. Rillig. 2012. Fungal superhighways: Do common mycorrhizal networks enhance below ground communication? Trends in Plant Science 17: 633?637. Bidartondo, M. I. 2005. The evolutionary ecology of mycoheterotrophy. New Phytologist 167: 335?352. Callaway, R. M., D. Cipollini, K. Barto, G. C. Thelen, S. G. Hallett, D. Prati, K. Stinson, and J. Klironomos. 2008. Novel weapons: invasive plant suppresses fungal mutualists in America but not in its native Europe. Ecology 89: 1043?1055. Carey, E. V., M. J. Marler, and R. M. Callaway. 2004. Mycorrhizae transfer carbon from a native grass to an invasive weed: evidence from stable isotopes and physiology. Plant Ecology 172: 133?141. Duhamel, M., R. Pel, A. Ooms, H. B?cking, J. Jansa, J. Ellers, N. M. van Straalen, T. Wouda, P. Vandenkoornhuyse, and E. T. Kiers. 2013. Do fungivores trigger the transfer of protective metabolites from host plants to arbuscular mycorrhizal hyphae? Ecology 94: 2019?2029. Gorzelak, M. A., A. K. Asay, B. J. Pickles, and S. W. Simard. 2015. Inter-plant communication through mycorrhizal networks mediates complex adaptive behaviour in plant communities. AoB Plants 7. Lerat, S., R. Gauci, J. G. Catford, H. Vierheilig, Y. Pich?, and L. Lapointe. 2002. C14 transfer between the spring ephemeral Erythronium americanum and sugar maple saplings via arbuscular mycorrhizal fungi in natural stands. Oecologia 132: 181?187. Moola, F. M. and L. Vasseur. 2004. Recovery of late-seral vascular plants in a chronosequence of postclearcut forest stands in coastal Nova Scotia, Canada. Plant Ecology 172: 183?197. Querejeta, J. F., L.M. Egerton-Warburton, and M.F. Allen. 2003. Direct nocturnal water transfer from oaks to their mycorrhizal symbionts during severe soil drying. Oecologia 134: 55?64. Rhoades, D. F. 1983. Responses of alder and willow to attack by tent caterpillars and webworms: evidence for pheromonal sensitivity of willows. In: Plant Resistance to Insects: Based on a symposium sponsored by the ACS Division of Pesticide Chemistry at the 183rd Meeting of the American Chemical Society, Las Vegas, Nevada, March 28?April 2, 1982, ed. Paul Hedin. ACS Symposium Series number 208. pp. 55?68. American Chemical Society. Ryan, F. 2002. Darwin's Blind Spot: Evolution Beyond Natural Selection. Boston: Houghton Mifflin Harcourt. Simard, S. W., K. J. Beiler, M. A. Bingham, J. R. Deslippe, L. J. Philip, and F. P. Teste. 2012. Mycorrhizal networks: mechanisms, ecology and modelling. Fungal Biology Reviews 26: 39?60. Smith, S. E. and D. J. Read. 2008. Mycorrhizal Symbiosis, third edition. Cambridge, Massachusetts: Academic Press. Song, Y. Y., R. S. Zeng, J. F. Xu, J. Li , X. Shen, and W. G. Yihdego. 2010. Interplant communication of tomato plants through underground common mycorrhizal networks. PLoS ONE 5(10): e13324. Song, Y. Y., S. W. Simard, A. Carroll, W. W. Mohn, and R. S. Zeng. 2015. Defoliation of interior Douglas-fir elicits carbon transfer and stress signalling to ponderosa pine neighbors through ectomycorrhizal networks. Scientific Reports 5, Article no. 8495. Winther, J. L. and W. E. Friedman. 2007. Arbuscular mycor rhizal symbionts in Botr ychium (Ophioglossaceae). American Journal of Botany 94: 1248?1255. David Yih is president of the Connecticut Botanical Society. "},{"has_event_date":0,"type":"arnoldia","title":"Ernest Jesse Palmer and Charles Sprague Sargent: A Serendipitous Relationship","article_sequence":2,"start_page":12,"end_page":24,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25626","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14e8526.jpg","volume":75,"issue_number":2,"year":2017,"series":null,"season":null,"authors":"Palmer, Theodore W.","article_content":"Ernest Jesse Palmer and Charles Sprague Sargent: A Serendipitous Relationship Theodore W. Palmer Ernest Jesse Palmer (1875?1962) was a highly accomplished plant collector, botanical taxonomist, and naturalist. He was employed by the Arnold Arboretum from 1913 to 1948 and, during the latter part of his tenure, lived with his family in the old farm house on Centre Street (near the Arboretum's current greenhouses). In this article, Palmer's son, Theodore W. Palmer, explains the unlikely circumstances that made this exceptional career possible. E rnest Jesse Palmer was born in England but came to the United States when he was three years old, his family first settling in west central Missouri. From early childhood Palmer had a strong interest in natural history, an interest that was encouraged by his parents who were surprisingly well informed on many subjects although they had no significant formal education. However, when his father lost the ability to work in his midfifties, Palmer's free time and chances for his own formal education were severely restricted as he became the main source of support for his family at the age of eleven. After the family moved in 1891 to the lead and zinc mining boom town of Webb City, in southwestern Missouri, he concentrated his natural history interest on the Carboniferous age marine fossils which were abundant in the mine tailing piles. Charles Sprague Sargent (1841?1927), founding director of the Arnold Arboretum, was the person mainly responsible for directing Palmer's career to botany. Palmer and Sargent first became aware of each other through the botanist Benjamin Franklin Bush (1858?1937). Bush had started collecting plants for Sargent and the Arnold Arboretum around 1899, after several years of doing the same for the Missouri Botanical Garden. In his obituary of Bush, Palmer wrote that in 1900 he read Bush's 1894 paper: In the introduction the author requested that teachers, horticulturists, or anyone interested in the woody plants of the state, send specimens to him for identification, for the purpose of bringing about a better knowledge of the flora of the state and of the distribution of the trees and shrubs. Finding many plants unknown to me, I decided to take advantage of this opportunity, and mailed a small package of twigs and leaves to him. Bush identified botanical specimens for Palmer over several months, and arranged to visit the next year in April. He stayed with the Palmers for nearly a week and left a supply of driers for the crude plant press Palmer had constructed. After that, Bush visited Palmer repeatedly in Webb City. Palmer sent his first specimens, some hawthorn (Crataegus) fruits, to Sargent in November 1901. His first attempt at shipment failed, however, as Sargent wrote to say that \"many of the paper packages inside the bag broke open in transit so that the fruit was mixed. In future the fruit should be put in separate cloth bags.\" No doubt numbers of inept people sent such packages to the Arboretum. But Palmer corrected the matter with a second box of specimens and received the following praise from Sargent: I have your letter of the 18th and also your box of specimens. These are excellent and you have been very successful in drying them, for Crataegus is one of the most difficult of all genera to handle for the herbarium. I hope you realize how greatly I am indebted to you for your assistance in this investigation. When the spring opens I COURTESY OF THE AUTHOR COURTESY OF PRAIRIE MOON NURSERY ARNOLD ARBORETUM ARCHIVES Ernest Jesse Palmer (behind horse) in 1895, ready to deliver groceries with a horse-drawn wagon, one of many jobs he worked as a way of supporting his family. Bush's poppy mallow (Callirhoe bushii), a herbaceous perennial native to Missouri, Kansas, Arkansas, and Oklahoma, was named in honor of botanist Benjamin Franklin Bush. Charles Sprague Sargent in the Arboretum collections, photographed in 1907. HARVARD UNIVERSITY HERBARIA 14 Arnoldia 75\/2 ? November 2017 In his book Trees and Shrubs: Illustrations of new and little known ligneous plants (Volume 2, page 67), Sargent notes that he had botanized with Palmer near Webb City on October 2, 1901, finding the type specimen of Crataegus aspera (now known as C. pruinosa). Seen here is one of the specimens of C. aspera collected by Sargent that day. BIODIVERSITY HERITAGE LIBRARY, MISSOURI BOTANICAL GARDEN Ernest Jesse Palmer 15 and Shrubs, Volume I, p. 57, where he wrote: This handsome tree of the Crusgalli Group, one of the largest and most symmetrical of American Thorns, is named for its discover, Mr. E. J. Palmer, of Webb City, who has carefully collected and studied Crataegus in southwestern Missouri, where the genus is represented by a large number of interesting forms, of which several are still undescribed. Perhaps Palmer's early success was not particularly remarkable--Crataegus in southwest Missouri was widespread and varied. On the other hand, Palmer was able rather soon to distinguish the interesting from the commonplace, and therein lies the genius of any good collector. Thus began a correspondence that was eventually to change Palmer's life profoundly. During the next several years, until 1907, the letters were sporadic. These were the years during which Palmer was building his taxonomic skills as well as pursuing his many other interests in natural history--all while also supporting his family with multiple jobs, starting with delivering heavy loads with his father's horse and wagon in his teenage years to being chief bookkeeper for a local oil company. A drawing of Crataegus palmeri (now known as C. reverchonii var. palmIt is obvious that Sargent valued eri) from Sargent's Trees and Shrubs: illustrations of new or little known Palmer's work on his favorite subligneous plants, Plate 29. ject, Crataegus. Palmer was eager to hope that you will be full of enthusiasm please all his correspondents as can be seen in his correspondence with the many small-time and ready for a new campaign, for the field collectors with whom he exchanged specimens. about Webb City is by no means exhausted He did no less for this great man who showed yet. I shall write you later just what it an interest in him. He worked to master Crais desirable to look after in the spring. taegus as well as anyone could. By 1908 one (November 22, 1901) of his more colorful acquaintances, the ReverBy 1903, Sargent had published three new end John Davis of Hannibal, Missouri, wrote, species from Palmer's herbarium specimens: \"Mr. Bush says you know the several species Crataegus palmeri, C. lanuginosa, and C. [of Crataegus] now almost by heart; can sight them afar off, and call them all by name.\" Thus speciosa, naming the first for Palmer in Trees COURTESY OF THE AUTHOR 16 Arnoldia 75\/2 ? November 2017 The Palmer's house in Webb City, Missouri, which was designed and built by hand from scrap lumber in the mid1890s by E. J. Palmer and his father. Palmer's sister and mother are on the front porch. Palmer had at this early date achieved a local reputation with this confusing genus. Is it, then, surprising that Sargent wanted to hire Palmer to collect for him? On March 25, 1907, Sargent wrote: I now enclose a list of the Crataegus material collected by you or me in your region which has not yet been described. Much of it is incomplete and I very much hope that you may be able to do more work this year on these numbers. Won't you kindly write me what the prospect is and whether there is any chance of your extending your work beyond the immediate neighborhood of Webb City and Carthage? Eureka Springs in Arkansas is evidently a good field as we collected flowers there a good many years ago but have never had any fruit. Joplin, I take to be a rich field and unexplored; indeed I fancy there is no place in southwestern Missouri that you could visit without finding new forms. But Palmer's answer was negative at this time because of his need to support his family. Sargent visited Webb City in the autumn of 1907 and stayed in the Palmer family home, as noted Dutch botanist Hugo de Vries had done in 1904. (The family, who lived in impecunious circumstances, took pride in these visits.) On the visit itself, there is no information. Palmer was then 32 years old, knew his area well, and no doubt Sargent saw a fair sampling of the surrounding countryside. Early in 1908 Sargent again wrote asking Palmer to collect professionally: I have been very much impressed with the carefulness with which you have made collections and observations of Crataegus and your grasp of the genus. I wish you could devote more time to collecting and studying the southern Missouri. Would it be possible for you to make an arrangement with your employers by which you could get off a month in the spring and a month or six weeks in the autumn for this purpose? THE STATE HISTORICAL SOCIETY OF MISSOURI Ernest Jesse Palmer 17 WEBB CITY N OE L A Rand-McNally map, circa 1888, of southwestern Missouri; note the many railway lines, but no roads, that are shown. If this could be done I should be very glad to pay you the salary you now get and, of course, all your expenses. Will you think this over and let me know if it is not possible to make some such arrangement? I should like to think that the Arboretum could employ you continuously for a year or two, at least, in this sort of work, and I was rather in hopes that Professor Trelease [of the Missouri Botanical Garden] would join me in such a scheme but he does not see his way clear to doing so, and single-handed I do not see how we can manage it.... (February 18, 1908) Again Palmer felt unable to do so; he wrote that he couldn't keep his position (as accountant with the Waters-Pierce Oil Company) and still take off so much time. A Growing Partnership By 1910 the relationship between the two men was such that Sargent could ask Palmer to collect very specific specimens, although Palmer would have to go to some trouble to do so. For example, on January 14, 1910, Sargent wrote: I believe you know that curious Hickory which grows at Noel and for which Bush suggested the name of subvillosa. It is one of the minima set but has smooth bark and very broad leaflets. If you know where to find the trees, would it be possible for you to run down to Noel now and get us some winter branchlets showing winter buds, etc. Noel is nearly fifty miles from Webb City and this was before automobiles were common. There was the train, of course. Palmer said in later years that he could not have collected so widely had not the railway and electric railway afforded access to places some distance from Webb City. By this means he could go to the towns of Alba, Joplin, Duenweg, Carthage, and Galena. (In 1918 the line was extended to include Baxter Springs and Pitcher.) The fare was twenty cents round trip to anywhere NANCY ROSE 18 Arnoldia 75\/2 ? November 2017 In 1902, near Webb City, Missouri, E. J. Palmer collected the seeds from which this cockspur hawthorn (Crataegus crus-galli, accession 12079-A, photo from October 2017) was grown. within a fifty mile radius, and Palmer would take his bicycle along to give him mobility once he reached his destination. In the case under discussion, Palmer did secure the hickory specimen. He would, throughout the remaining years as a collector, go far out of his way to obtain a desired plant for Sargent. In 1911, Sargent urged Palmer again to collect professionally and for a more extended period than the snatched weekends and his annual two week vacation, which had been the only available time previously. The letters speak for themselves: You have a salary, I understand, of $50 a month in your present position and I understood from you that you were rather anxious to get a couple of months next summer to look after some changes and improvements in your house. It has occurred to me that possibly you might be willing to give up your present position for six months, beginning March 1st, and devote March, April, May, June, September, and October to collecting for the Arboretum, taking July and August for your own work. We could offer you the same salary that you have now and of course pay all your traveling expenses. If you can see your way to accepting such an offer, I should want you to go to Texas to look after some Crataegi there in March and then gradually work northward. There are indications in southern Missouri of a large number of still undescribed species of Crataegus of which we have incomplete material. I should be very glad to get these doubtful species cleared up and generally to get as much work done in southern Missouri, Arkansas, and eastern Texas as time will permit. I am very anxious to get this work done and I don't know any one so well fitted to do it as yourself, so I hope that I shall get a favorable answer to this letter. (December 20, 1911) ARNOLD ARBORETUM ARCHIVES Ernest Jesse Palmer 19 A sketch by E. J. Palmer of hawthorn (Crataegus) leaves and fruit. And, only a few weeks later: ... My idea is that the proposed field work would take practically all the time from March 1st to December and January unless you wanted, as I had supposed, a couple of months at home in the summer for your work. I hope this arrangement can be made for I feel very strongly the importance of it and that you are the best man for the work. If the proposition I made you does not appear satisfactory, let me know just what you want. I think if possible we ought to decide pretty soon because my idea would be for you to go to Texas early in March and it will take some time to properly lay out the campaign, etc. In any case do not let this thing fall through if there is any possible way of preventing it. (January 11, 1912) Another negative answer from Palmer prompted this reply by Sargent: I am very much disappointed at your inability to devote the summer to botany and I am still in hopes that some arrangement may be made. The situation is this. We have indications of a large number of new species of trees and shrubs in southern Missouri, Arkansas, and eastern Texas. To collect these intelligently it is necessary that the collector should be in a position to visit the localities in the spring and autumn. Bush cannot be depended on for this as he is often tied up by his business. The result of this in the past has been that we have hundreds of incomplete specimens. I do not see why, if you want to take up botany, you cannot have summer employment for three or four years at least, and possibly make as much or more than you do now with the possibility that this work might lead to something better in the future. Of course if you collected for the Arboretum you could at the same time do what Bush 20 Arnoldia 75\/2 ? November 2017 A New Genus BRENT BAKER, ARKANSAS NATURAL HERITAGE COMMISSION In 1913 Palmer collected an unknown plant in Jasper County, of which Bush wrote, \"Do not send out any of No. 3921, and if you have any more of it, please send me all of it until I can get it described ... This is a new genus to be known as Geolobium minimum ... I have found many new species, but never a new genus, and a genus so peculiarly situated, with no known relative, no known family to receive it.\" The name Geolobium was dropped in favor of Geocarpon and the plant was described by Kenneth Kent Mackenzie (to whom Bush apparently sent it for identification) in 1914. Mackenzie placed it in Aizoaceae, the fig-marigold or ice plant family, but it was changed much later to Caryophyllaceae, the pink family. Incidentally, it appeared that at least some of Palmer's botanical employers were annoyed that the description of the plant had fallen to Mackenzie, who was a lawyer and amateur botanist allied with the New York Botanical Garden. His connection with Palmer was an indirect one through Bush. While the United States had been a gold mine of new genera for a few hundred years, by 1913 a new genus was fairly uncommon (among phanerogams, certainly) and to have one get away to another institution hurt more than a little. Geocarpon minimum is a tiny (less than 2 inches [5 centimeters] tall), fleshy plant that grows natively in a limited number of sites in Missouri, Arkansas, Louisiana, and Texas. did, make sets for yourself and sell them with the understanding that they were not to be distributed until the plants had been named. I wish you would give this matter serious consideration for I think it is of the utmost importance to American botany that these collections should be made and I feel sure that the chances for you in advancements in the next few years would be better than what they would seem to me to be if you remain as a clerk in Webb City. Perhaps you can at least tell me what sort of a proposition you might want if you are not satisfied with the one I have made. (February 1, 1912) Bush wrote Palmer in April 1912, after a visit, that he wanted to show Palmer over the collecting ground because it was likely that he would succeed Bush in the field. Bush also wrote, \"You know more about them [Crataegus] than I do or anyone else does for that matter.\" However, Palmer did not yet accept an offer from Sargent and in May Sargent wrote, \"I am still most anxious that you should be connected with the Arboretum as a collector and I shall never lose an opportunity of suggesting to you the desirability of your Palmer's saxifrage (Saxifraga palmeri, syn. Micranthes palmeri) is one of several plants named in honor of E. J. Palmer, this species by his colleague accepting my propositions.\" Again, early in 1913, Sargent Benjamin Franklin Bush. This small herbaceous plant grows on rocky sites in open woodlands in Arkansas and Oklahoma. wrote: remain here for a few days at the ArboreI am in a position now to arrange permanent botanical employment for you for a tum.... The sooner you come the better, period of at least three years at a salary for the offer, if you accept it, means the which will be in advance of what you are beginning of work in the early spring ...\" now getting. I think this is a proposition (January 20, 1913) which you ought not to hesitate to accept Palmer's quick negative reply elicited this even if you are tied up in mining ventures, response from Sargent: which you can certainly turn over to some I regret extremely that you do not see your one else. Before deciding either to accept way to taking up botany as a profession. or refuse this offer I want to talk it over I wanted to open a way for you to have a fully with you and I suggest that you come really distinguished career and one that to Boston at once, at my expense, and HENDRIX COLLEGE HERBARIUM Ernest Jesse Palmer 21 22 Arnoldia 75\/2 ? November 2017 ARNOLD ARBORETUM ARCHIVES would have paid you better than your present occupation. Mining properties are terribly uncertain and in nine cases out of ten, I am afraid, they lead to loss and disappointment. I had supposed that if you saw your way to accepting my proposition you would make your headquarters in St. Louis, and that your mother and sister would move there where I believe they would find life as comfortable and pleasant as in Webb City. I cannot tell you how disappointed I am at your decision. (January 28, 1913) The farm house at 1090 Centre Street is seen in a black-and-white photograph from the early 1900s and a contemporary view, photographed around 2000 by the author's childhood friend, Henry Alfred Anderson. In a letter from Palmer to Sargent on March 31, 1913, Palmer again declined Sargent's request, though this time he left open the possibility of future work, writing: I am situated just as I was when I wrote you a few weeks ago, and I do not see how it would be possible for me to get away from here at present. I could not leave or dispose of my mining interests at this time without sacrificing all that I have put into them. However, I expect to know within a few months whether I shall realize anything on my investment or not, and it has been chiefly on this account that I have delayed making any change in my plans for the past year. As I am situated in Webb City, with my mother and sister here and some property to look after, I can scarcely see how I could manage to be away altogether. However, I expect to make a change of some sort shortly, but not likely before fall. If there is still an opening in the line you suggest at that time I might be in a position to take advantage of it, but could not possibly do so just now. Ernest Jesse Palmer 23 Moving to Boston It was not until after his mother, Anna Windle Palmer (born 1841) died on April 30, 1920, that Palmer felt free to move to Boston. Finally at 11:00 p.m., April 4, 1921, a cold and rainy night, at age 45, Palmer arrived in Boston. He had never visited an eastern city before and knew only Sargent in the whole of Boston. The next day, Palmer called on Sargent before noon. (That evening he wrote his sister that the $1.50 he paid for a simple breakfast put him \"on the road to bankruptcy.\") The small Arboretum staff (all paid quite modestly) had been looking for an apartment for Palmer when he arrived. The best that they had found was a two-room suite upstairs in the house of the superintendent of the grounds, Christian Van der Voet, some distance from a place to eat. At the Arboretum, Palmer worked six and a half days a week. That half day, Sunday morning, was the most important time since Sargent arrived usually before 8 a.m. (and therefore so did Palmer). They enjoyed working together for a relatively uninterrupted few hours. Generally no one else was there on Sunday to consult the great professor. Every day Palmer rose in the morning about 5:30 a.m., spending about an hour before going out for breakfast. Then he worked until 12:30 or 1:00 p.m. when he had a small lunch at a food stand near the Arboretum. After dinner at a restaurant, he took an evening's walk, bought the newspaper, and went home to read. When the weather per mitted, on his after noon off, Palmer explored Boston on foot. (Besides relishing the exercise, Palmer deplored the high price of a trolley ride: ten cents.) Through the years Palmer continued to go on plant collecting expeditions as well as working in the Arboretum's collections and herbarium, and writing extensively on plants and other natural history topics, including the Native American artifacts he collected on the grounds. Sargent's death in March 1927 was very distressing for Palmer. He had lost a friend and mentor and then, as of April 30 that year, Ernest Henry Wilson terminated Palmer's job, purportedly as a cost cutting measure (by Wilson's estimation the Arboretum was $120,000-- nearly 1.7 million in today's dollars--in debt at the time of Sargent's death). Harvard botanist Oakes Ames (1874?1950) was appointed Supervisor of the Arboretum in June 1927, and COURTESY OF THE AUTHOR His resistance was fading, however, and he wrote to Sargent on June 20, 1913, \"While it would be more advantageous to me in my present financial circumstances to retain my present position until spring I scarcely feel that I should put you off any longer if you feel that the work to be done this year is urgent, and I am to undertake it.\" He finally began working as a collector for the Arnold Arboretum and Missouri Botanical Garden later that year. The death of his father on September 17, 1911, after a long illness, probably made this easier. E. J. Palmer, at age 78, putting plants in a herbarium press while out botanizing, which he did regularly long after retirement. 24 Arnoldia 75\/2 ? November 2017 ARNOLD ARBORETUM ARCHIVES servants often also enjoy proving that they can get along well on their own in primitive situations. Palmer officially retired from the Arboretum around midsummer in 1947, but returned to work in the herbarium from September of that year until early summer 1948. Palmer and his family then moved back to the family home in Webb City, Missouri. He continued to work on botany and other natural history interests seven days a week, despite declining eyesight. On his death bed, he dictated the last few words of a botanical article about his beloved Ozark forest to his wife, Elizabeth. He died hours later, on February 25, 1962. Palmer published more than 100 botanical papers in his career. Academic writing was not his only forte, though-- in 1958, Elizabeth gathered seventy-six of her husband's poems, the result of his lifelong love of poetry, and published them in a volume titled Gathered Leaves, Green, Gold and Sere. Acknowledgements E. J. Palmer reading his poetry book, Gathered Leaves, Green, Gold and Sere, at home in Missouri. Wilson was appointed Keeper at about the same time. Ames and Palmer had already established a good relationship and with Ames now as Supervisor, Palmer was rehired by 1928. In 1930, at age 55, Palmer married Elizabeth McDougal, a bacteriologist at the Massachusetts State Laboratory, which is located to the east of the Arboretum. They had three children--my brother, sister, and me. Ames arranged for the family to live in the house at 1090 Centre Street, owned by Harvard University but adjacent to and controlled by the Arboretum. My father's stories when I was a young child left no doubt that he and Sargent had greatly enjoyed each other's company. Throughout his career Sargent managed to enlist a number of people down on their luck in a variety of ways to become part of his \"band of brothers\" and sisters. During Palmer's collecting years, Sargent repeatedly made arrangements to spend a few days with him in the field. Wealthy men like Sargent who enjoy the luxury of many My wife, Laramie Palmer, began writing a Master's thesis on my father for the University of Kansas in 1970. I have freely used her extensive but unfinished manuscript for this article and in a biography I hope to publish commercially in which she is acknowledged as co-author. She should be recognized as co-author here, but I wanted to be able to say \"my father.\" The many letters between Sargent and Palmer are in the Arnold Arboretum Archives. Palmer kept copies also and my family donated them to the State Historical Society of Missouri. I have received permission from both institutions to quote from the letters. Bibliography Bush, B. F. 1895. A list of the trees, shrubs, and vines of Missouri. Missouri Horticultural Society Report, pp. 353?393. Kobuski, C. E. Ernest Jesse Palmer, 1875?1962. Journal of the Arnold Arboretum 43: 351?358. Palmer, E. J. 1937. Benjamin Franklin Bush. American Midland Naturalist 18(3): 1?6. Sargent, C. S. 1905. Trees and Shrubs: Illustrations of new and little known ligneous plants. Boston and New York: Houghton Mifflin Company. Theodore W. Palmer is Emeritus Professor of Mathematics at the University of Oregon and one of the founders of Mount Pisgah Arboretum near Eugene, Oregon. "},{"has_event_date":0,"type":"arnoldia","title":"From Fungi to Forests: The Tale of Tropical Tree Diversity","article_sequence":3,"start_page":25,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25626","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14e8526.jpg","volume":75,"issue_number":2,"year":2017,"series":null,"season":null,"authors":"Krishnadas, Meghna","article_content":"From Fungi to Forests: The Tale of Tropical Tree Diversity Meghna Krishnadas T in close proximity? It appears that the answer might partly lie in plant pathogens (MullerLandau 2014). How pathogens regulate the diversity of trees is a remarkable ecological tale. But, to better understand this story, we need to make a short trip down a path of scientific hypotheses. Many ideas have been posited to explain the conundrum of how multiple species coexist at relatively small spatial scales to generate diversity. Broadly, diversity can stem from mechanisms classified as stochastic or deterministic. Simply put, stochastic mechanisms are happenstance. ALL PHOTOS BY ARUN KUMAR AND MEGHNA KRISHNADAS hink biodiversity and a tropical rainforest might come to mind. Wet tropical forests, the most species-rich ecosystems on earth, are estimated to support nearly half of all known terrestrial species. Trees are a particularly diverse group of organisms in wet tropical forests and some regions like the Amazon can host over 1,000 different tree species in a single hectare of forest. Ecologists have long pondered the mechanisms underlying such astounding variety (Wright 2002). How do so many organisms of such similar, albeit simple, requirements coexist in such diversity Forest fragmentation creates edges, and edge effects occur when differences in habitat conditions in relation to distance from edge also change ecological communities. 26 Arnoldia 75\/2 ? November 2017 The Natural History of Natural Enemies Predators and pathogens regulating population numbers of their prey or host is a well-known concept in a variety of ecosystems. In rivers, lakes, seas, or forests, predators keep consumer numbers in check. As with larger animals, predators of seeds and seedlings also exert a topdown effect on plant populations, and wet tropical forests are home to a variety of insects and fungi that kill seeds and seedlings. The forest understory provides especially conGerminating seed of Drypetes oblongifolia, an evergreen tree native to the ducive settings for fungi. Thriving in Western Ghats in southern India. the wet, dark forest floor, soil fungi infect seeds and seedlings, thus regulating their numbers. In a surprising twist, however, by keeping down the numbers of their hosts, plant predators allow other species to persist, thus promoting diversity of plant communities. Pathogens act in a manner termed \"negatively density dependent.\" Negative density dependence just means that an individual plant is more likely to be infected by its pathogen when in the vicinity of its own kind, i.e., other individuals of the same or closely related plant species (Freckleton and Lewis 2006). Just as human diseases spread more Seeds being matched to seedlings in the greenhouse. when humans crowd together, Would a seed arrive at a spot? Would a seedincreasing individuals of a single plant species ling be accidentally killed by a falling branch cultivates the soil for the very pathogens that or underfoot a roving deer? In contrast, deterkill the plant's seeds and seedlings. So, when ministic mechanisms, as the term suggests, are a species becomes numerous, its individuals processes that govern species' survival in preare more likely to die, allowing less competidictable ways, such as availability of necessary tive species to persist, leading to a more diverse resources or susceptibility to pests, pathogens, community. and predators (Freckleton and Lewis 2006). Back in 1971, negative density dependence The interaction between plants and their as a mechanism for diversity in multispecies consumers and plants and pathogens has been communities was proposed independently by suggested previously as an important driver Janzen and Connell, for tropical rainforests of plant diversity. Only recently, however, and coral reefs, respectively (Janzen 1971). The have the links between pathogens and plant Janzen-Connell (J-C) hypothesis states that diversity been demonstrated through empiridiversity of a community is maintained in part cal experiments. by the parasites and predators that cull young Tropical Tree Diversity 27 insects and mammalian seed predators, although evidence was mounting that fungal pathogens were also crucial agents of negative density dependence. Notably, no study had explicitly tested whether densitydependent culling by seed and seedling predators actually increased diversity of the plant community. Then in 2014, in a seasonally wet forest in Belize, researchers demonstrated how the plant community changed if the action of insects and fungi was experimentally inhibited using insecticide and fungicide (Bagchi et al. 2014). Without insects, Roads and other linear clearings fragment forests and the resulting edge the community of recruiting seedeffects alter the recruitment dynamics of plant communities. lings was markedly different from the naturally regenerating community. Without fungi, the diversity of seedlings dropped sharply in relation to natural regeneration. Clearly, pathogens drove diversity of the tree community in this neotropical forest. But would insects and fungi play the same role in other closed-canopy forests? Moreover, insect and fungal communities vary with factors like light and moisture--factors that also affect plants directly. How would pathogen impacts on plant diversity and composition change with different habitat conditions brought about from different light and moisture levels? A hard edge where forest abruptly transitions to a field; habitat conditions are Importantly, with environmental very different for seedling establishment here. conditions changing rapidly because progeny, especially where host numbers are of human actions, how would maintenance of high. Specifically, mortality of seeds and seedplant diversity via pathogens change in humanlings of a species will be higher with increasing altered forests (Swinfield et al. 2012)? numbers of that species in a neighborhood (such What Happens When Humans as close to seed-producing adult trees), opening Alter Forests? up these spaces for other species to occupy, thus promoting diversity. We live today in a human-dominated planet. For A recent meta-analysis--an analysis of the food, fuel, and other natural resources, humans net outcome from multiple studies conducted have deforested much of the earth, breaking in different ecosystems--found that overall once-large and contiguous forest into smaller empirical evidence supported the J-C hypothparcels, a process known as forest fragmentaesis (Comita et al. 2014). However, the J-C tion. One of the greatest threats to biodiversity, hypothesis was largely tested in the context of forest fragmentation sets in motion a range of 28 Arnoldia 75\/2 ? November 2017 ecological processes that alter the dynamics of species' survival in the remnant forests. Long-term research in experimentally and naturally fragmented forests have found that fragments often lose species in predictable ways (Laurence et al. 2011). Edge effects, or altered habitat conditions at forest edges, are strongly associated with changes in fragmented tree communities. At edges and in smaller fragments, slow-growing, dense-wooded, shade-tolerant tree species are lost over time, being replaced by fast-growing, light- Human activities like agriculture are another major cause of forest fragmenloving species. However, while pat- tation and change in ecological communities terns of species' losses are clear, the underlying mechanisms are less well understood (Didham et al. 2012). Hitherto, edge effects have been primarily examined as changes to abiotic conditions--alteration in light, moisture, wind speed, etc., as we move from the forest edge to interior. Because species differ in their ability to survive in different levels of these resources, changes to light or moisture are assumed to alter species survival at different distances from edges. As a consequence, the plant community changes at edges compared to interior forest. Such changes in the tree community have important consequences for ecosysA typical human-dominated tropical landscape where forests exist in a matrix tem functions such as carbon stor- of farms, roads, and human habitations age or nutrient cycling (Chapin et al. 2000). Hence, grasping the mechanisms drivshade-tolerant species are able to come in. ing community-wide changes to trees could Without abundant light, the light-loving species do not have the resources to grow fast and improve management and inform restoration are outcompeted by slower-growing species. of fragmented forests for tree diversity and ecosystem function. Events unfold differently in forest edges. So why are edges dominated by light-wooded, Even after the canopy forms, shade-tolerant early successional tree species? Let us suppose species seem unable to establish at edges. It is that edges behave like giant forest gaps. Lightargued that high light at edges disadvantages loving, fast-growing species often colonize and slow-growing, shade-tolerant species, and fastdominate gaps by exploiting the high resource growing species outcompete them. As a consequence, edges and small fragments, which conditions. But, once the canopy is established are subject to edge effects, remain dominated in a gap, light availability reduces and more Tropical Tree Diversity 29 by light-loving species while shade-tolerant species are unable to regain a foothold. However, it is also possible that this \"arrested succession\" is happening because of changes to pathogen activity. Light-loving species tend to be more susceptible to pathogens than shade-tolerant species, although there are exceptions. While lightloving species might initially increase in areas of high light, their numbers should start coming down when pathogens build up around them over time. Thus, in edges of older fragments, the seedlings of abundant light-loving species should suffer higher mortality from pathogens, opening up that space for shade-tolerant species. However, warmer, drier conditions at edges might reduce pathogen activity, thus diluting the mechanism that prevents one of few species from becoming super abundant. Alternatively, the benefits of high light per se help overcome losses to pathogens for all species. In this case, no species will be much affected by pathogens at edges and controlling pathogen activity would not improve survival of shade-tolerant versus shade-intolerant species. Plant?Pathogen Interactions in Fragmented Forest In a fragmented, human-altered forest, I examined whether and how the influence of pathogens during seedling recruitment (establishment and survival of seedlings) varied with distance to edges. The research site was within the Western Ghats Biodiversity Hotspot in Karnataka state, India. To test whether light alone or a combination of light and pathogens regulated seedling recruitment, I set up groups of seedling plots at increasing distances from the forest edge. Each group consisted of two seed traps and five seedling plots. In each A one-by-one-meter plot where seedlings have been tagged and identified. 30 Arnoldia 75\/2 ? November 2017 Plots were demarcated and labeled at their diagonal ends using PVC pipes, colored ribbons, and a location code. A tagged seedling of Litsea floribunda infected by leaf fungi, resulting in a nectrotic spot. group, one plot each was sprayed with fungicide, insecticide, fungicide plus insecticide, and water, and one plot was retained as control without any spraying. I set up 145 such groups at 15 locations, three groups each at distances of 0, 25, 50, and 100 meters (0, 82, 164, and 328 feet) from the edge, totaling 730 seedling plots. I applied pesticide treatments from November 2015 through November 2016. During this time, seeds falling into the seed traps were recorded twice a month. I conducted censuses for new recruits twice during the year: once at the end of the dry season and then at the end of the wet season after peak recruitment occurred. Preliminary results indicate that seedling diversity reduces when plots are sprayed with fungicide, but only as we move into interior forest. Similarly, turnover of species between seeds that arrive at a spot and seedlings that establish is lowered with fungicides, but only in interior forest. Importantly, the density-dependent effect of fungi and insects appears to be at play only in interior forest. Hence, it appears that the lower diversity of seedlings in plots with pesticides are likely due to a loss of pathogen-mediated mortality of seeds and young seedlings as we move towards the forest edge. Clearly, edge effects are changing some interactions between plants and their pathogens, which in turn appears to be affecting the diversity of the plant community. The Future of Diversity New recruits (seedlings) were censused and tagged twice during the experiments. Uncovering the mechanisms driving diversity remains a fascinating quest. You know you have stumbled upon a rich question when every answer opens up more questions. But, as we slowly piece together bits of the diversity puzzle, we are also changing natural systems at an unprecedented scale. Labeling the \"Anthropocene\" as a valid geological epoch awaits scientific consensus, but few can miss the ubiquitous influence of Tropical Tree Diversity 31 humans on Earth (Corlett 2015). Human actions have wiped out entire species, introduced new plants and animals to places where they were unlikely to reach, changed species' numbers in relation to one another, and altered biological communities in a blink of evolutionary time. Unless we apply some serious course correction, today's biodiversity might stand a bleak chance for tomorrow. Even if we set aside areas of land and water for other species, much of Earth will likely continue to be occupied by Homo sapiens in the near future. One hopes that by understanding the subtle processes that generate diversity, we can better manage the spaces that we share with other species, both for biodiversity and its contribution to human needs. Bibliography Bagchi, R. et al. 2014. Pathogens and insect herbivores drive rainforest plant diversity and composition. Nature 506: 85?88. Corlett, R. T. 2015. The Anthropocene concept in ecology and conservation. Trends in Ecology and Evolution 1?6. doi:10.1016\/j.tree.2014.10.007 Didham, R. K., V. Kapos, and R. M. Ewers. 2012. Rethinking the conceptual foundations of habitat fragmentation research. Oikos 121: 161?170. Freckleton, R. P. and O. T. Lewis. 2006. Pathogens, density dependence and the coexistence of tropical trees. Proceedings of the Royal Society, Biological Sciences 273: 2909?2916. Janzen, D. H. 1971. Seed predation by animals. Annual Review of Ecology and Systematics 2: 465?492. Laurance, W. F. et al. 2011. The fate of Amazonian forest fragments: A 32-year investigation. Biological Conservation 144: 56?67. Muller-Landau, H. C. 2014. Ecology: plant diversity rooted in pathogens. Nature 506: 44?45. Swinfield, T., O. T. Lewis, R. Bagchi, and R. P. Freckleton. 2012. Consequences of changing rainfall for fungal pathogen-induced mortality in tropical tree seedlings. Ecology and Evolution 2: 1408?1413. Chapin, F. S. et al. 2000. Consequences of changing biodiversity. Nature 405: 234?242. Wright, S. J. 2002. Plant diversity in tropical forests: a review of mechanisms of species coexistence. Oecologia 130: 1?14. Comita, L. S. et al. 2014. Testing predictions of the Janzen-Connell hypothesis: a meta-analysis of experimental evidence for distance- and density-dependent seed and seedling survival. Journal of Ecology 102: 845?856. Meghna Krishnadas is a graduate student in the Yale School of Forestry and Environmental Studies and the 2016 recipient of the Arnold Arboretum's Ashton Award for research in Asian tropical forest biology. 36673667 U.S. POSTAL SERVICESTATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION(Required by 39 U.S.C. 3685) 1. Publication Title: Arnoldia. 2. Publication No: 0004?2633. 3. Filing Date: September 29, 2017. 4. Issue Frequency: Quarterly. 5. No. of Issues Published Annually: 4. 6. Annual Subscription Price: $20.00 domestic; $25.00 foreign. 7. Complete Mailing Address of Known Office of Publication: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 8. Complete Mailing Address of Headquarters of General Business Office of Publisher: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 9. Full Names and Complete Mailing Address of Publisher, Editor, and Managing Editor: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500, publisher; Nancy Rose, Arnold Arboretum, 125 Arborway, Boston, MA 02130?3500, editor. 10. Owner: The Arnold Arboretum of Harvard University, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 11. Known Bondholders, Mortgagees, and Other Security Holders Owning or Holding 1 Percent or More of Total Amount of Bonds, Mortgages, or Other Securities: none. 12. The purpose, function, and nonprofit status of this organization and the exempt status for federal income tax purposes have not changed during the preceding 12 months. 13. Publication Name: Arnoldia. 14. Issue Date for Circulation Data Below: August 17, 2017. 15. Extent and Nature of Circulation. a. Total No. Copies. Average No. Copies Each Issue During Preceding 12 Months: 1775. Actual No. Copies of Single Issue Published Nearest to Filing Date: 1600. b. 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I certify that all information furnished on this form is true and complete. Nancy Rose, Editor. "},{"has_event_date":0,"type":"arnoldia","title":"Fraxinus bungeana: An Ash of a Different Color","article_sequence":4,"start_page":32,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25628","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14e896f.jpg","volume":75,"issue_number":2,"year":2017,"series":null,"season":null,"authors":"Aiello, Anthony S.","article_content":"Fraxinus bungeana: An Ash of a Different Color Anthony S. Aiello F raxinus bungeana, Bunge ash, is a plant that challenges one's preconception of a genus because, unlike other ashes, this Chinese native is a shrub with showy flowers instead of a tree with inconspicuous flowers. It belongs to a group known as the flowering ashes, which includes the better-known F. ornus along with the Asian F. sieboldii and F. chinensis. Its flowers and form are more reminiscent of its oleaceous cousins, Syringa (lilac) and Chionanthus (fringetree), than they are of familiar members of its own genus. This species has been cultivated in botanic gardens since the late 1800s, but has never significantly jumped the fence into general horticulture. Fraxinus bungeana is native to north-central China, where it grows in dry sandy soils and rock crevices. It was first collected by intrepid Russian plant collector Alexander von Bunge in 1831 on one of his explorations of Siberia, Mongolia, and Beijing. It was subsequently given its species name by botanist Alphonse de Candolle in 1844. Additional herbarium records appear throughout the latter half of the eighteenth century, but the first known seed introduction into North America was to the Arnold Arboretum in April 1880 from the Mus?um National d'Histoire Naturelle, in Paris. A second collection was received by the Arboretum in January 1882, from Emil Bretschneider, a physician and botanist who explored the plains and mountains surrounding Beijing. Remarkably, one plant (accession 14625*A) from this collection is still alive, located in the ash collection on the east side of Bussey Hill. As of July 2017, this tree had a height of 12 feet (3.7 meters) and spread of 16 feet (4.9 meters). The next wild collections after Bretschneider's were those of the USDA's explorer, Frank Meyer, who collected seeds in late 1907, again in and around Beijing. As far as can be determined, none of Meyer's plants remain alive. Meyer aptly described it as, \"an ash growing in rocky situations and on steep mountain sides. Attains, apparently, no great size. May be of use as a foresting plant in semiarid regions.\" In light of the spread of emerald ash borer, and as part of recent efforts to increase the diversity of ashes in the United States, collections of F. bungeana seeds were made in 2009 by Kang Wang, Beijing Botanical Garden, in Liaoning, Hebei, and rural Beijing municipality, and the following year by Kang Wang, Michael Dosmann (Arnold Arboretum), and I, near Beijing, as part of the 2010 North America-China Plant Exploration Consortium expedition (collections NACPEC10-039 and NACPEC10-042). Bunge's ash is a large shrub, growing 6 to 15 feet (1.8 to 4.6 meters) tall. The plants from which we collected in China in 2010 were 3 to 6 feet (0.9 to 1.8 meters) tall. As mentioned above, it has small, terminal flower panicles with small white fine-petaled corollas that resemble those of fringetree. Alfred Rehder, in his Manual of Cultivated Trees and Shrubs, described it as a \"distinct species, handsome in bloom.\" This year I noticed seed production for the first time on one plant at the Morris Arboretum. Fall foliage color is at best similar to forsythia, with purple overtones fading to yellow. As with other ashes, Bunge ash prefers full sun, and seems adaptable to a range of soil pH. It clearly can tolerate dry conditions but also thrives in rich soils. It is likely cold hardy through USDA Zone 5 (average annual minimum temperature -10 to -20?F [-23.3 to -28.9?C]). Anthony S. Aiello is the Gayle E. Maloney Director of Horticulture and Curator at the Morris Arboretum of the University of Pennsylvania in Philadelphia. Cultivar description: Fraxinus bungeana `Sunflash' Growing plants from seed can result in a great amount of variation, providing the opportunity to select for vigor and health and, occasionally, to find novel plants. This was the case in September 2014 when I first noticed a variegated plant among a group of seedlings from the NACPEC10-042 Fraxinus bungeana collection (Morris Arboretum accession number 2010-213*A). This plant, which has not flowered yet, has a growth rate and habit similar to others collected in 2010, but differs in having leaves that are patterned with a mixture of green, yellow, and cream. This variegation has been consistent since it was first observed and persists throughout the season. We are currently working on propagating this individual for further distribution. Fraxinus bungeana `Sunflash' "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23456","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14eab28.jpg","title":"2017-75-2","volume":75,"issue_number":2,"year":2017,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Plant Exudates and Amber: Their Origin and Uses","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25624","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14e8128.jpg","volume":75,"issue_number":1,"year":2017,"series":null,"season":null,"authors":"Lambert, Joseph B.; Santiago-Blay, Jorge A.","article_content":"Plant Exudates and Amber: Their Origin and Uses Jorge A. Santiago-Blay and Joseph B. Lambert P some other plant pathology. In other instances, such as in typical underground roots, exudate production appears to be part of the typical metabolism of healthy plants that helps stabilize the soil and foster interactions with other organisms around the roots. Different plant tissue types and organs can produce exudates. We have collected resins and gums from the above ground portions of plants, or shoots, as well as from the generally below ground portion of plants, or roots. Root exudation has been known for decades and is respon- ALL PHOTOGRAPHS BY JORGE A. SANTIAGO-BLAY UNLESS OTHERWISE NOTED REPRODUCED WITH PERMISSION OF AMERICAN SCIENTIST lants produce and export many different molecules out of their cellular and organismal confines. Some of those chemicals become so abundant that we can see or smell them. The most visible materials oozed by many plants are called \"exudates.\" What are plant exudates? Generally, exudates are carbon-rich materials that many plants produce and release externally. When exudates are produced, they are often sticky to human touch. Such plant chemicals can be the visible expression of attack by bacteria, fungi, herbivores, or Resinous exudates on a conifer. Prolific white, resinous exudation is seen on a tumorlike growth on the trunk of a white pine (Pinus strobus) at the Arnold Arboretum. Blobs of white resin on a relatively young shoot of a Japanese black pine (Pinus thunbergii, AA accession 11371-O). Plant Exudates and Amber 3 A slab of Great Basin bristlecone pine (Pinus longaeva) right out of the microwave oven showing extruded (and very hot!) resinous exudates. Microwave heating experiments were performed at the Laboratory of Tree-Ring Research, University of Arizona, Tucson. sible for many of the fascinating relationships in the interface of plant roots and soil microorganisms known as the rhizosphere. Collecting and Analyzing Plant Exudates After receiving collecting permission (if needed), we spend days walking the grounds of botanical gardens and arboreta, or do field work elsewhere. Exudates are easily collected directly from the trees with no harm to the plant and leaving no doubt about their botanical identity. Occasionally we use more forceful methods, such as carefully microwaving wood slabs to extract the exudates, then letting them resolidify. Once the material is collected, we place it in a small plastic zip-top bag. An additional, external bag is used to hold a paper label containing the collection data. If needed, we let the exudate dry slowly in an oven and, once dried, the materials are ready for subsequent analyses. In other instances, generous collaborators send us materials for chemical analyses. Carbon-13 solid state Nuclear Magnetic Resonance spectroscopy (ssNMR) is a stateof-the-art research tool that generates spectra (or chemical signatures) of materials, including plant exudates and amber or greatly fossilized plant resin. The analyses, which use a tiny amount (as little as 50 to 100 milligrams, approximately the volume of a new eraser on a school pencil) of the exudate, are non-destructive. They are performed at Northwestern University (in Evanston, Illinois), one of a few research laboratories in the world with carbon-13 ssNMR capabilities. At times, we observe plants that evidently have produced exudates but the amounts are insufficient for our analyses. Solid exudates are pulverized manually and undergo two sets of carbon-13 ssNMR analyses: normal decoupling, which gathers signals for all carbon atoms, and interrupted decoupling, which, among others, obtains signals from carbons lacking the attached hydrogens. Just like in spectra used in the health-allied sciences, different regions of the spectra provide valuable information (see Figure 1 on page 4). In the case of NMR, the peaks represent different atoms and reflect their molecular environment. The height of the peaks largely represents rela- 4 Arnoldia 75\/1 ? August 2017 (A) Interrupted Decoupling ? only carbons with strong C-H interactions analyzed Normal Decoupling ? all carbons analyzed C=O bonds as in carboxyls C=C bonds as in alkenes C-O bonds as in sugars C-C bonds as in alkanes 220200180160140120100 80 60 40 20 0 f1 (ppm) (B) Interrupted Decoupling ? only carbons with strong C-H interactions analyzed Normal Decoupling ? all carbons analyzed C=O bonds as in carboxyls C=C bonds as in alkenes C-O bonds as in sugars C-C bonds as in alkanes 220200180160140120100 80 60 40 20 0 (ppm) (C) Interrupted Decoupling ? only carbons with strong C-H interactions analyzed Normal Decoupling ? all carbons analyzed C=O bonds as in carboxyls C=C bonds as in alkenes C-O bonds as in sugars C-C bonds as in alkanes 220200180160140120100 80 60 40 20 0 (ppm) Figure 1. Chemical identity of peaks on a C-13 ssNMR spectra. Panel (A) is a resin, panel (B) is a gum, and panel (C) is a kino (a type of phenolic, often found in Eucalyptus). In all panels, the upper result uses interrupted decoupling, which eliminates peaks representing C-H single bonds. The lower result uses normal decoupling in which all carbon-to-atom bonds are represented. tive abundance of those atoms. The position of the peak along the horizontal axis (parts per million [ppm]) is the resonance frequency characteristic of the atom and its molecular neighborhood. This position is an indication of the chemical identity of the peak as compared to an external molecular reference. In carbon-13 ssNMR, peaks in the 0?80 ppm region are singly bonded carbon atoms (-C-C-), or alkanes; signals within the 80?100 ppm region are single bonded carbon atoms with electron-withdrawing neighbors, in particular, oxygen (C-O), as found in carbohydrates, such as sugars. Currently, we have analyzed over 1,800 exudates of all types, including amber, representing most of the major plant groups worldwide. However, a lot more samples still need to be acquired and analyzed. Types of Plant Exudates Using NMR, we have determined that there are three major types of plant exudates: resins, gums, and phenolics. Resins are made from terpene molecules. The basic molecular unit of terpenes is a five-carbon molecule, known as isoprene (see Figure 2 on page 6). When freshly produced, many resins are sticky and smell like Christmas trees or incense. Resins are insoluble in water and thus do not dissolve during rains. As time passes and the resins begin to \"mature,\" many of their original chemical constituents evaporate. The materials remaining behind in the resin blob form chemical bonds, a process known as polymerization, and the blob begins to harden. With the passage of millennia, the resinous material becomes greatly polymerized and Plant Exudates and Amber 5 Not On the Collection List JOSEPH O'BRIEN, USDA FOREST SERVICE, BUGWOOD. ORG Not everything that looks like an exudate is an exudate. Some living organisms, particularly fungi, can resemble the kinds of plant exudates we collect. In other instances, the watery--and often foul smelling--material that decomposing portions of plants produce can also resemble exudates. As you may guess, we do not collect those! Clockwise: Some exudate mimics include a cedar-apple rust (Gymnosporangium juniperi-virginianae) fungal fruiting body on Juniperus virginiana; an unidentified fungus growing on a Pinus hwangshanensis (AA accession 68-76-F)--note its superficial similarity to the yellowish color of some resins; a Polyporus fungus on Quercus palustris (AA accession 805-87-A); an exudateresembling, foul smelling material resulting from decomposition by fungi and bacteria on a cut Cornus kousa (AA accession 524-49-D) branch. 6 Arnoldia 75\/1 ? August 2017 CHIP CLARK, NATIONAL MUSEUM OF NATURAL HISTORY, SMITHSONIAN INSTITUTION CH3 CH2 H2C Figure 2. An isoprene molecule, the building block of resins. CH2OH H C HO CHIP CLARK, NATIONAL MUSEUM OF NATURAL HISTORY, SMITHSONIAN INSTITUTION Close-up of resinous flow on the trunk of a pine (Pinus). C H OH C O OH H C C H OH H Figure 3. Model of a glucose, an example of a simple sugar molecule. Chemically linked sugar molecules make up carbohydrates. The carbon bound by two oxygen atoms (arrow) is known as anomeric carbon and is characteristic of sugars. Exudated carbohydrates are known as gums. Latex exudate emanating from a Euphorbia tirucalli stem. evolves into the robust gemstone called amber, produced only by specific plant species. Conifers such as pines (Pinus), firs (Abies), spruces (Picea), larches (Larix), and some other familiar cone-bearing trees in northern latitudes tend to produce resinous exudates. Many angiosperms (flowering plants) also produce resins. The term \"latex\" refers to milky-looking exudates produced by numerous flowering plants, including those in the euphorbia or spurge family (Euphorbiaceae). Latexes can be dangerous to touch, causing dermatitis or other damage, especially to the eyes. Interestingly, all latexes we have examined thus far are resins in suspension. A second type of exudates is known as gums. Gums are large carbohydrates consisting of myriad sugar molecules linked together chemically (see Figure 3 above). Gums do not CHIP CLARK, NATIONAL MUSEUM OF NATURAL HISTORY, SMITHSONIAN INSTITUTION Plant Exudates and Amber 7 Gum produced by a Yoshino cherry (Prunus ? yedoensis) growing near the Tidal Basin in Washington, D.C. Reddish phenolic exudates are visible on the trunk of this Eucalyptus sideroxylon. tend to smell because of their low volatility stemming from their high molecular weight. When freshly produced, many gums are spongy to touch because of their high water content. Thus, freshly produced gums dissolve easily during rains. If somehow gums manage to survive and dry out, they can then be very hard to dissolve. However, as far as we are aware, gums are not known to survive millions of years as amber does. Gum exudates tend to be produced by flowering plants; fruit trees in the genus Prunus, including cherries, plums, peaches, and almonds, commonly produce gums. The third major type of exudates is known as phenolics. Phenolics are chemically related to terpenes but form unsaturated ring compounds known as aromatics because of their often-pleasant odor. When freshly produced, phenolics tend to be watery and reddish brown, and lack the strong smell of resins. If they survive dissolution, phenolics tend to form brittle solids. As with gums, we are not aware of phenolics that have survived deep time. Phenolics tend to be common in Eucalyptus and related plants. Combinations of these major types of exudates, such as gum resins, as well as several other minor kinds of exudates are also known. Uses of Plant Exudates In addition to their generally beautiful colors, pleasant aroma, and light weight, resins are water insoluble. These properties make resins, including amber, coveted natural products. Some uses of resins, including amber, include: ceremonial and artistic, as construction materials, ingestive, and, of course, as objects of science because they provide windows into past worlds. Ceremonial and artistic uses Amber, that is, greatly polymerized resin, has been used for ceremonial purposes as well as for objects of trade, jewelry, sculptures, and many other items. Although highly valued in the market, amber varies greatly in color and translucency, from white to black and from translucent PATRICK R. CRAIG CHIP CLARK, NATIONAL MUSEUM OF NATURAL HISTORY, SMITHSONIAN INSTITUTION 8 Arnoldia 75\/1 ? August 2017 Earrings made from Columbian copal were treated in an autoclave, which applies heat and pressure, resulting in a color change from yellow to green. NSAA, WIKIMEDIA COMMONS PATRICK R. CRAIG An assortment of typical yellowish amber specimens showing the wide range in color and translucency. PATRICK R. CRAIG A group of typical Baltic amber specimens shows varying color. Specimens of rare Dominican blue amber from the personal collection of Patrick R. Craig. Retsina is a Greek wine traditionally flavored with pine resin. to opaque. Because of this variability, color and translucency on their own are generally not good diagnostic traits for identifying amber. On the other hand, copal (less polymerized resin) and modern resins are still used in some areas of Mexico and Central America for artistic and ceremonial purposes, prized because they smell of incense. Next time you encounter a pine, fir, or spruce tree, look carefully at its bark and you may be able to see some exudate blobs or \"teardrops.\" Pick one of them up and smell it! Pine resin has been used in the preparation of rosin, which is applied to the hairs of bows used to play string instruments such Plant Exudates and Amber 9 Tertiary amber Mesozoic amber Is It Amber or Copal? Amber is greatly fossilized resin. This resinous fossilized material has been found in numerous localities worldwide. The oldest amber has been dated as early as the Carboniferous period, over 300 million years ago. Often, forests whose trees produced resins that eventually became amber tended to be located close to sea level at the time of production. Partially polymerized resin is known as copal, a Nahuatl or Aztec word that means incense. At times, we have seen the term \"semi-amber\" used instead of copal. We recommend avoiding the term \"semi-amber\" because it suggests the material is older than it really is. Although it can be difficult to distinguish copal from resin, a straightforward preliminary way to distinguish between the two is by using a drop of organic chemical such as 95% ethanol or acetone (the solvent used in most nail polish removers). Take a drop of the chemical and place it in a portion of the test sample that has little or no value to the owner. Then touch the wetted portion with the finger. If it feels sticky, the test sample likely is copal; if it does not feel sticky, likely it is amber. We have examined a number of alleged amber samples that turned out to be copal, some of which were in the collections of respectable museums. When finding \"amber\" specimens of potential scientific value, we recommend testing them by physicochemical means, such as nuclear magnetic resonance spectroscopy (NMR) or others, to gain more confidence on the specimen's true nature. 10 Arnoldia 75\/1 ? August 2017 as the violin (rosin makes the hairs just sticky enough to grip the strings and create sound). ROBERT MAYER Construction materials The metallic transatlantic cable that connected the Old and New Worlds telegraphically during the second half of the nineteenth century was insulated by gutta percha, the resinous exudate of Palaquium gutta, a tropical Southeast Asian tree. The modern aviation and aerospace industry uses human-made, lightweight and strong, synthetic resins and phenolics in building airplanes. Ingestive An old and interesting use of resins is in the preparation of retsina, a Greek wine that is flavored with a little bit of pine resin (typically from Aleppo pine, Pinus halapensis). Gums are also sometimes eaten; in places where the legumiCollecting Competition nous Acacia trees produce copious Interestingly, sometimes birds, such as the types of woodpeckers quantities of gums, these exudates commonly called sapsuckers (genus Sphyrapicus), compete with are used as survival foods when us as they also feed on exudates and leave characteristic holes on other food is scarce. Although it has the surface of some trees. Other birds and some insects are known been alleged that amber has healing to use exudates for nest construction. and other medicinal properties, we are not aware of scientific studies using a double-blind protocol that demonstrate any medicinal properties of amber. A yellow-bellied sapsucker (Sphyrapicus varius) perches on a conifer branch that displays the typical holes created by this and other sapsucker species. Science For reasons that are not known, some forests in the past appear to have produced copious amounts of resins. Although these exudates may have attracted some organisms and repelled others, once small organisms such as insects landed on the sticky material it was difficult to detach from it. When subsequent resin flows covered the specimen it was protected from the action of decomposing organisms and the environment, allowing it to be preserved for a longer time. Subsequent polymerization of the resin preserved a fraction of the resin-entombed organisms, which, when found, now have great value to scientists. Amber encased plant and animal specimens have contributed insights in a number of scientific fields. Amber specimens that contain larger, rarely found organisms (e.g., scorpions, amphibians, lizards, birds) are of great interest and may command great sums of money. How- PATRICK R. CRAIG Plant Exudates and Amber 11 PATRICK R. CRAIG A drosophilid fly trapped in amber. unknown botanical origin, like the resin from the Java Sea wreck, with those in our database. With that information, we were able to suggest that the plant whose resins were harvested back in the thirteenth century was from the botanical family Dipterocarpaceae, and perhaps specifically the genus Shorea. Having an idea of the botanical provenance of archeological artifacts enriches our knowledge of how our predecessors used plants. In this case, research tells us that aromatic resins were an important commodity at the time and were often imported into China for use in Buddhist rituals as well as medicines, lacquers, and perfumes. We will continue to collect and analyze plant exudates from around the world, including amber and copal, as well as materials associated with anthropological artifacts, adding knowledge for future researchers to use. References Kosmowska-Ceranowicz, B. 2015 Infrared spectra atlas of fossil resins, subfossil resins and selected imitations of amber. In: ATLAS, Infrared Spectra of the World's Resins, Holotype Characteristics. pp. 3?213. Warszawa, Polska: Polska Akademia Nauk Muzeum Ziemi w Warszawie. Wood fibers encased in amber. ever, buyer beware, as there are unscrupulous sellers willing to make money from objects that are not genuine amber. Ongoing Research Goals Ultimately, we seek answers to questions because we are curious about nature. Sometimes, our results can help answer a question. For example, along with several other colleagues, including Dr. Lisa Niziolek from the Field Museum of Natural History in Chicago, we answered the question: In what plant family was the tree that produced the blocks of resin found in a thirteenth century shipwreck excavated from the Java Sea? Our studies of many plant exudates have generated a large database of their NMR profiles. When we study a sample of unknown botanical provenance, that database allows us to compare the samples of Lambert, J. B., C. E. Shawl, G. O. Poinar, Jr., and J. A. Santiago-Blay. 1999. Classification of modern resins by solid nuclear magnetic resonance spectroscopy. Bioorganic Chemistry 27: 409?433. Lambert, J. B., Y. Wu, and J. A. Santiago-Blay. 2005. Taxonomic and chemical relationships revealed by nuclear magnetic resonance spectra of plant exudates. Journal of Natural Products 68: 635?648. Lambert, J. B., Y. Wu, and J. A. Santiago-Blay. 2002. Modern and ancient resins from Africa and the Americas. In: Archaeological Chemistry. Materials, Methods, and Meaning. Chapter 6, pp. 64?83. Symposium Series No. 831. K. A. Jakes (Editor). American Chemical Society. Washington, District of Columbia. Lambert, J. B., M. A. Kozminski, C. A. Fahlstrom, and J. A. Santiago-Blay. 2007. Proton nuclear magnetic resonance characterization of resins from the family Pinaceae. Journal of Natural Products 70(2): 188?195. Lambert, J. B., M. A. Kozminski, and J. A. Santiago-Blay. 2007. Distinctions among conifer exudates by proton magnetic resonance spectroscopy. Journal of Natural Products 70(8): 1283?1294. 12 Arnoldia 75\/1 ? August 2017 Lambert, J. B., Y. Wu, and M. A. Kozminski, and J. A. Santiago-Blay. 2007. Characterization of Eucalyptus and chemically related exudates by nuclear magnetic resonance spectroscopy. Australian Journal of Chemistry 60: 862?870. Lambert, J. B., J. A. Santiago-Blay, and K. B. Anderson. 2008. Chemical signatures of fossilized resins and recent plant exudates. Mini Review. Angewandte Chemie (International Edition) 47: 9608?9616. Also published in German, with the following bibliographic information: Chemischer Fingerabdruck von fossilen Harzen und rezenten Pflanzenexsudaten. Angewandte Chemie 120: 9750?9760. Lambert, J. B, E. A. Heckenbach, Y. Wu, and J. A. SantiagoBlay. 2010. Characterization of plant exudates by principal component and cluster analysis with nuclear magnetic resonance variables. Journal of Natural Products 73(10): 1643?1648. Lambert, J. B., C. Y.?H. Tsai, M. C. Shah, A. E. Hurtley, and J. A. Santiago-Blay. 2012. Distinguishing amber classes by proton magnetic resonance spectroscopy. Archaeometry 54(2): 332?348. SUZANNE C. SHAFFER Lambert, J. B., C. L. Johnson, E. W. Donnelly, E. A. Heckenbach, Y. Wu, and J. A. SantiagoBlay. 2013. Exudates from the asterids: characterization by nuclear magnetic resonance spectroscopy. Life: The Excitement of Biology 1(1): 17?52. SUZANNE C. SHAFFER Lambert, J. B., E. R. Heckenbach A. E. Hurtley, Y. Wu, and J. A. Santiago-Blay. 2009. Nuclear magnetic resonance spectroscopic characterization of legume exudates. Journal of Natural Products 72: 1028?1035. On the lookout even during vacation, author Jorge A. Santiago-Blay (left) noticed resinous exudates on several lodgepole pines (Pinus contorta) in Yellowstone National Park, including one partially debarked, possibly by American bison (Bison bison) (right). Note the copious exudate production (yellowish color) on the debarked portion of the trunk. Plant Exudates and Amber 13 Lambert, J. B., E. W. Donnelly, E. A. Heckenbach, C. L. Johnson, M. A. Kozminski, Y. Wu, and J. A. Santiago-Blay. 2013. Molecular classification of t he nat u r a l e xuda t e s of t he r os i ds . Phytochemistry 94: 171?183. Lambert, J. B., A. J. Levy, J. A. Santiago-Blay, and Y. Wu. 2013. NMR characterization of Indonesian amber. Life: The Excitement of Biology 1(3): 136?155. Lambert, J. B., J. A. Santiago-Blay, Y. Wu, and A. J. Levy. 2014. Examination of amber and related materials by nuclear magnetic resonance s p e c t r o s c o p y. M a g n e t i c R e s o n a n c e i n Chemistry (Special Issue on NMR in Cultural Heritage) 53: 2?8. Lambert, J. B., J. A. Santiago-Blay, R. Rodr?guez Ramos, Y. Wu, and A. J. Levy. 2014. Fossilized, semifossilized, and modern resins from the Caribbean Basin and surrounding regions for possible preColumbian Trans-Caribbean cultural contacts. Life: The Excitement of Biology 2(4): 180?209. Lambert, J. B., C. L. Johnson, A. J. Levy, J. A. SantiagoBlay, and Y. Wu. 2015. Molecular classification of exudates from the monocots, magnoliids, and basal eudicots. Life: The Excitement of Biology 3(2): 083?117. Lambert, J. B., J. A. Santiago-Blay, Y. Wu, and A. Levy. 2016. The structure of stantienite. Bulletin for the History of Chemistry 40(2): 86?94. Lambert, J. B., C. L. Johnson, T. M. Nguyen, Y. Wu, and J. A. Santiago-Blay. 2016. Ferns, cycads, Ginkgo, and Gnetophytes: Nuclear Magnetic Resonance characterization of exudates from exotic plant sources. Life: The Excitement of Biology 4(3): 215?232. https:\/\/blaypublishers. files.wordpress.com\/2016\/11\/lambert-et-al2016-leb-43215-2321.pdf Lambert, J. B. Y. Wu, and J. A. Santiago-Blay. 2016. Highresolution solid-state NMR spectroscopy of cultural organic materials. In: Webb, G. Modern Magnetic Resonance. Second Edition. Springer. Lambert, J. B., A. J. Levy, L. C. Niziolek, G. M. Fienman, P. J. Gayford, J. A. Santiago-Blay, and Y. Wu. 2017. The resinous cargo of a Java Sea shipwreck. Archaeometry. (A paper authored by M. Donahue describing this research was published in The Smithsonian Insider on May 15, 2017. http:\/\/insider.si.edu\/2017\/05\/resin-shipwreckhints-trade-routes-botany-ancient-asia\/ .) Langenheim, J. H. 2003. Plant Resins: Chemistry, Evolution, Ecology, and Ethnobotany. Portland, Oregon: Timber Press. Mills, J. S. and R. White, R. 1994. The Organic Chemistry of Museum Objects. Second Edition. Oxford, England: Butterworth-Heineman. Nussinovich, A. 2010. Plant Gum Exudates of the World: Sources, Distribution, Properties, and Applications. Boca Raton, Florida: CRC Press. Rodr?guez Ramos, R., J. Pag?n Jim?nez, J. A. SantiagoBlay, J. B. Lambert, and P. R. Craig. 2013. Some indigenous uses of plants in pre-Columbian Puerto Rico. Life: The Excitement of Biology 1(1): 83?90. Santiago-Blay, J. A., R. L. Hoffman, J. B. Lambert, and Y. Wu. 2003. Cylindroiulus truncorum (Silvestri): a new milliped for Virginia (USA), with natural history observations (Julida: Julidae). Banisteria 20: 62?66. Santiago-Blay, J. A. and J. B. Lambert. 2007. Amber's botanical origins uncovered. American Scientist 95: 150?157. (Reprinted with permission as Aux sources de l'ambre. Pour la Science [French version of Scientific American] June 2007. 356: 70?75. Abstracted by David M. Kondo in the Winter 2007 issue of Gems and Gemology 43: 395.) Santiago-Blay, J. A. and J. B. Lambert. 2010. Legumes and their exudates. Aridus (Bulletin of the Desert Legume Program of the Boyce Thompson Southwestern Arboretum and the University of Arizona) 22(1): 1, 4, 6. Santiago-Blay, J. A. and J. B. Lambert. 2010. Desert plants and their exudates. Desert Plants 26 (1): 1, 3?8. Santiago-Blay, J. A., J. B. Lambert, and P. P. Creasman. 2011. Expanded applications of dendrochronology collections: Collect and save exudates. TreeRing Research 67(1): 67?68. V?vra, N. 2015. Mineral names used for fossil resins, subfossil resins and similar materials. In: ATLAS. Infrared Spectra of the World's Resins ? Holotype Characteristics. pp. 215?280. Warszawa, Polska: Polska Akademia Nauk Muzeum Ziemi w Warszawie. Dedication Author Jorge A. Santiago-Blay dedicates this paper to his mother, ?ngeles Blay S?lomons, who in the early 1980s suggested to him that he pursue the study of \"las resinitas\" (the little resins) as she used to call exudates. Her memory always lives with him. Jorge A. Santiago-Blay is a Resident Research Associate in the Department of Paleobiology at the Smithsonian Institution's National Museum of Natural History in Washington, D.C. (blayj@si.edu). Joseph B. Lambert is Research Professor of Chemistry at Trinity University in San Antonio, Texas , and Clare Hamilton Hall Professor of Chemistry Emeritus, Northwestern University, Evanston, Illinois (jlambert@northwestern.edu). "},{"has_event_date":0,"type":"arnoldia","title":"Other Order: Sound Walk for an Urban Wild","article_sequence":2,"start_page":14,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25623","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14ebb6f.jpg","volume":75,"issue_number":1,"year":2017,"series":null,"season":null,"authors":"Rueb, Teri; Del Tredici, Peter","article_content":"Other Order: Sound Walk for an Urban Wild Peter Del Tredici and Teri Rueb I n urban areas, vegetation that is not planted or maintained by people--including both native and non-native species--typically dominates many different habitats including river and stream banks, highway verges, vacant building lots, infrastructure edges, chain-link fence lines, and random pavement cracks. For most cities, the amount of spontaneous vegetation they support varies inversely with their economic prosperity; cities that have lost the most population and jobs show the highest levels of land abandonment and volunteer plant growth (Del Tredici 2010a, b; Burkholder 2012). How city dwellers respond to the presence of spontaneous vegetation in their midst is influenced by personal preferences as well as by cultural norms. In many European cities, residents' feelings about spontaneous vegetation is divided--some welcome it as a manifestation of unrestrained urban nature while others see it as an indicator of dereliction that should be removed. Such responses have led to the categorization of urban residents as either \"nature lovers\" (a.k.a. wilderness enthusiasts) or \"neat freaks\" (a.k.a. urban devotees). Interestingly, the percentages of people in these categories can vary dramatically from one city to the next in the same country (Keil 2005; Rink 2005; Weber et al. 2014). In an effort to promote a wider appreciation and acceptance of \"urban wilds,\" urban ecologists have recently been attempting to calculate the value of the ecosystem services provided by spontaneous vegetation, especially in cities where the population is shrinking and the amount of vacant land is expanding (Pataki et al. 2011; Burkholder 2012; Robinson and Lundholm 2012). On the positive side, this vegetation contributes to increasing the ecological functionality of the city in terms of storm water management, temperature reduction, carbon sequestration, soil development, and total biodiversity (Kowarik and K?rner 2005; Carroll 2011). Harder to quantify, but nevertheless important, are the opportunities for social, cultural, educational, and nature experiences that spontaneous vegetation provides across the wide array of cultural contexts and sites that characterize most cities (Pfeiffer and Voeks 2008; Daniel et al. 2012; Jorgensen and Keenan 2012). The aesthetics of spontaneous vegetation are usually considered negative given that much of it is perceived as ugly or messy (i.e., lacking ornamental characteristics or possessing an unkempt appearance), and its presence in the landscape is justifiably viewed as projecting an image of neglect (Nassauer and Raskin 2014). In the arena of public health, many people see spontaneous vegetation as providing habitat for animals that are vectors for a number of human pathogens and infectious diseases such as rats, mosquitoes, and ticks (Garvin et al. 2012; Gulachensik et al. 2016). Similarly, the large size that spontaneous urban vegetation can reach in the absence of maintenance is viewed as providing cover for potential criminal activity and thus a threat to public safety. To the extent that urban landscapes dominated by spontaneous urban vegetation are perceived as threatening, they fit within a concept of a \"wilderness\" that is defined as land that exists outside the bounds of human control (Hofmeister 2010; Jorgensen and Keenan 2012; Desimini 2015). In this article we will explore the history of the \"urban wilds\" construct as it developed in Boston, Massachusetts, from its introduction in the mid-1970s through today, and present a case study of one such site, Bussey Brook Meadow at the Arnold Arboretum, to illustrate how multi-faceted urban wilds can be creatively interpreted for the general public utilizing GPS (global positioning system)-based cell phone technology. ALL PHOTOS BY PETER DEL TREDICI UNLESS OTHERWISE INDICATED Other Order: Sound Walk 15 Urban Wilds in Boston Urban vegetation takes hold in cracks in a neglected swath of asphalt pavement. Pretty wildflowers or invasive weeds? Chicory (Cichorium intybus), yellow sweet clover (Melilotus officinalis), and spotted knapweed (Centaurea stoebe subsp. micranthos) bloom along a city street. Weedy trees take over an abandoned lot in Detroit. In the United States, the idea that unmanaged \"open space\" in cities could perform valuable ecological services was foreshadowed by a movement in the 1970s that categorized such sites as \"urban wilds\" (Tanner 1975; Desimini 2015). In 1976, the Boston Redevelopment Authority (BRA), a city planning agency, officially adopted the term when it issued an inventory of Boston's unimproved and unprotected natural areas under the title Boston Urban Wilds. The report--which was partially funded by a grant from the National Endowment for the Arts--was spearheaded by BRA landscape architect Elliot Rhodeside. It identified 143 parcels of land (2,000 total acres) of diverse sizes and ownerships that contained significant \"natural resource value\" but were threatened by on-going development pressure (BRA 1976). Most of the sites had histories of industrial, institutional, or residential use, some dating back to the nineteenth century. Rhodeside left the BRA shortly after the report was published and the work of advocacy, fundraising, and protection for the Boston Urban Wilds project passed to a non-profit organization, the Boston Natural Areas Fund (BNAF), founded in 1977 by Eugenie Beal (the head of the then newly formed Boston Conservation Commission) along with her future husband, John Blackwell. In its early days, BNAF was focused on trying to preserve and protect properties listed in the Urban Wilds report, but over time the emphasis of the organization shifted away from land acquisition to maintenance of already protected properties and coalition building with other nonprofit organizations around issues of public advocacy. In 1988, the Boston 16 Arnoldia 75\/1 ? August 2017 Boston Urban Wilds, a 1976 report from the Boston Redevelopment Authority, identified 143 land parcels in Boston with potential value as preserved natural areas. This copy of the report is in the Arnold Arboretum library. Parks Department officially took over management of the Urban Wilds program which, as of 2014, listed 39 properties in its inventory. The Parks Department currently provides maintenance and logistical support for those properties that are controlled by the city; other properties on the list receive varying levels of maintenance depending on the resources allocated by the organization that controls it (Bird 2014). The original 1976 BRA report described 143 sites that contained some significant \"natural resource value,\" including geological features (68 sites), coastal or fresh water wetlands (20 sites), shorelines (27 sites), or important vegetation (28 sites). It was clearly a simpler time when the meaning of the words nature and natural were not contested and the dichotomy between native and exotic species had yet to emerge as the divisive issue it is today. In the 1970s, urban wilds, regardless of their biological content or cultural history, were viewed as valuable antidotes to blighted, barren cityscapes. By the late-1990s, the original concept of an urban wild became subsumed under the rubric of ecological restoration. This reconceptualization of urban nature--essentially attempting to affix a \"native\" label on it--represented a dramatic reversal of fortune for the non-native organisms that found themselves reclassified as invasive species. Older, less value-driven terms to describe these plants, including weed, pest, naturalized species, garden escapee, volunteer, etc., fell by the wayside and with them an appreciation of their historic connection with the past land use of the site (Del Tredici 2010b). This privileging of native over non-native species has created problems for today's advocates of urban wilds because many of the sites they're striving to protect can no longer be \"restored\" to anything resembling their original ecological condition (Del Tredici 2010a, b; Carroll 2011). Similarly, the ways that some people use minimally maintained urban wilds, including drinking, doing drugs, having sex, painting graffiti, and camping out, has also caused problems for advocates because of complaints from abutting residents and other users. Like it or not, urban wilds are places where human behavior, like the plants and animals that occupy them, can sometimes be out of control (Keil 2005; Thompson 2012). Bussey Brook Meadow Bussey Brook Meadow of the Arnold Arboretum of Harvard University was listed as an urban wild in the 1976 BRA report. This 25-acre wetland has a documented history of land use going back 350 years when, in 1662, one of the first roads leading southwest out of the city of Boston was constructed along its western edge. This road opened the land up to farmers who drained portions of the property and moved the stream, Bussey Brook, that ran through the middle of it to the periphery. A hundred and forty years later, in 1802, another road was built along its eastern edge, which eventually developed into a railroad line that is still in operation today. Once a stable earthen berm was constructed for the rail line in 1873, Bussey Other Order: Sound Walk 17 Willows and cattails are among the moisture loving plants that thrive in Bussey Brook Meadow. Brook was effectively isolated from the larger, adjacent Stony Brook watershed that drains into the Charles River. Following this, the processes of fragmentation and filling of Bussey Brook Meadow accelerated dramatically, most notably with the installation in 1900 of a 9-foot-diameter, 3,600-foot-long high-level sewer line across the western edge of the property and the construction of an expanded Forest Hills train station to the north in 1909 (Arnold Arboretum). The Arnold Arboretum, through a land purchase from its Harvard University parent, acquired roughly half of the Bussey Brook Meadow parcel in 1919 and constructed Muddy Pond in the middle of the site, a wagon road following the track of the high-level sewer line, and a tree nursery on the site of former agricultural land. All of these activities over a period of seven years resulted in more wetland filling and disruption of surface drainage. In the 1950s, a privately owned esker on the site, composed mainly of sand and gravel, was excavated for construction purposes and replaced between 1955 and 1965 with a 5-acre landfill consisting of construction debris from the demolition of several Boston public schools. In 1971, after an unfortunate incident in which two neighborhood children from a nearby public housing project drowned, Muddy Pond in the center of the Arboretum's portion of the wetland was filled in (Arnold Arboretum). In 1982, the northern end of Bussey Brook Meadow underwent a major transformation when an expansion of the Forest Hills train station was initiated in order to make room for the new Orange Line subway station. As part of this project, a pathway was built along the base of the landfill that linked the new subway station to the South Street gate of the Arboretum. In 1996, through the determined efforts of 18 Arnoldia 75\/1 ? August 2017 JAY CONNOR Eugenie Beal and John Blackwell and the two non-profit organizations they co-founded (the Boston Natural Areas Fund and the Arboretum Park Conservancy), the deeds to the variously owned parcels of Bussey Brook Meadow were bundled together and added to the Arnold Arboretum's 1882 indenture, thereby achieving permanent protection for the entire 25-acre site. At the same time, grants from federal, state, and city agencies, together with funds generated by the Arboretum Park Conservancy, were used to construct granite entrance gates and upgrade the surface of the main pathway, now christened Blackwell Path. In 2011, the Arboretum, in keeping with its scientific research mission, approved a plan that called for turning Bussey Brook Meadow into a site for long-term environmental monitoring and research on urban ecology. While this meant that most of the land in Bussey Bussey Brook Meadow with Blackwell Path running through it. Brook Meadow would be left alone to follow its own ecological trajectory, the Arboretum made a commitment to manage portions of the site that were heavily used by visitors, including mowing the edges of Blackwell Path regularly, removing hazard trees that threatened public safety, and mowing several meadow areas annually to keep woody vegetation from taking over (Arnold Arboretum). Creating the Other Order Sound Walk Having developed a long-term strategy for the site, the Arboretum still had to contend with issues raised by discordant visitor perceptions of the land. Some saw Bussey Brook Meadow as a \"natural\" counterpoint to the well-maintained landscape of the Arnold Arboretum proper, while others saw it as a haven for invasive species that undermined the Arboretum's moral authority on matters of ecology. It was into this ARNOLD ARBORETUM ARCHIVES Other Order: Sound Walk 19 The 9-foot-diameter sewer line being installed in Bussey Brook Meadow in August 1900. Blackwell Path in Bussey Brook Meadow in summer. breech that the authors of this article stepped in 2012 with a proposal to interpret Bussey Brook Meadow for the general public using a GPSbased sound walk designed for use with a cell phone. The purpose of the proposed app was to illuminate the complex cultural history of Bussey Brook Meadow, to reveal the complex ecological interactions that are currently taking place on the site, and to show some of the ways the site was being used by the general public. The ultimate goal of the project was to try and change how people thought and felt about the site--to help them see that it was not just a chaotic collection of weeds but a dynamic, organized ecological system that reflected cultural A tree-of-heaven (Ailanthus altissima) grove on the slopes of the landfill in Bussey Brook Meadow. Ring counts of downed trees indicated that they established themselves on the site in 1965 or 1966, shortly after dumping stopped. values, past land-use history, and future ecological trajectories (Rueb and Del Tredici 2014). The Other Order app took two years to complete and involved a close collaboration between Del Tredici, who provided extensive verbal interpretation of the site in situ, and Rueb, who recorded this material and combined it with field recordings to create a soundscape designed for delivery via a downloadable mobile app. The app uses GPS to track visitors' movements and play the sounds at specific locations in Bussey Brook Meadow as they pass through them. In addition to Del Tredici's monologues on \"cosmopolitan\" vegetation, recordings included dozens of on-site conversations with 20 Arnoldia 75\/1 ? August 2017 Non-native wetland plants growing in Bussey Brook Meadow: common reed (Phragmites australis) in the background, yellow flag iris (Iris pseudacorus) in the middle, and reed canarygrass (Phalaris arundinacea) in the foreground. These species have sorted themselves out across a moisture gradient to form a functional urban wetland. various experts, stakeholders, Arboretum staff, and park visitors including urban ecologists, park advocacy groups, multi-generation urban farmers, landscape architects, dog walkers, commuters, and transient residents. Over twenty hours of ambient field recordings of the environment were also incorporated and used as inspiration for sound compositions evoking, for example, the material layers of the landfill, the interior sounds of the high level sewer pipeline, the leisure activity of past visitors mingled with those of the present, and the wildlife of the meadow. In some places one might find unexpected sounds such as light snoring tucked under a tree-of-heaven grove (Ailanthus altissima) near a concrete overhang, a flute mingling with sounds of laughter in a clearing in the center of the old Arboretum nursery, cows lowing and chickens clucking near the site of the former Bussey farmstead, and the sound of underwater gurgling as captured with hydrophones dropped into Bussey Brook. All of these sounds reference actual and imagined ways in which the meadow has been inhabited over time and the various materials and organisms that make up the complex social, biological, and physical matrix that is Bussey Brook Meadow. Sound regions are arranged throughout the meadow in a manner that allows for a complete experience should visitors constrain their movements solely to Blackwell Path, which takes about twenty minutes to traverse at a leisurely pace. However, additional sound regions Other Order: Sound Walk 21 A wetland in Bussey Brook Meadow consisting of common reed (Phragmites australis), yellow flag iris (Iris pseudacorus), and reed canarygrass (Phalaris arundinacea). are spread throughout the meadow, rewarding the more adventurous and patient with sounds that may be accessed only by leaving the trail behind and following informal footpaths, trails, and tunnels through the dense vegetation. Blackwell Path is an egalitarian corridor that connects the elegant environs of the formal Arboretum with the urban hubbub of the Forest Hills subway station and surrounding neighborhoods. Intervening into this path system represents a critique of conventional parks as much as an invitation to go off the beaten track and explore the wilds held within this \"urban wild.\" Bussey Brook Meadow is a particularly complex social site as it sits somewhere between a managed botanical garden on the one hand, and an interstitial zone where commuters, neighborhood residents, and tourists from all over the world mix with transient populations who are often staying for extended periods of time or returning each year with the milder seasons to regular encampments. A central concern of the work is to communicate a variety of perspectives on place as a means of critically engaging contested meanings, uses, and inhabitations of public sites. Voices in the work range from those of experts who tell us what to look for, what to hear, and what to value in this richly vegetated environment, to those that offer meandering impressions, personal histories, random thoughts, and idiosyncratic perceptions of a place. Animals, wind, weather, and water are equal voices in this mix. Through this blending of voices, the work draws upon the cosmopolitan botany of the site as a central metaphor and a means for JON HETMAN 22 Arnoldia 75\/1 ? August 2017 Peter Del Tredici and Teri Rueb recording conversations in Bussey Brook Meadow. asking probing questions about ownership, access, interpretation, and use of public parks and green spaces. Among the discoveries that stood out for the authors in developing the work and seeing its reception across various audiences was an awareness of the intensity of the experience as visitors were often torn between giving themselves over to immersion in the layered sounds emanating from their headphones and relating them to the complex sights, sounds, and social activities of the site itself. Frequently groups of people would walk together, taking their headphones off at regular intervals to exchange impressions and ask each other if they heard the same thing, and if it was \"in the headphones\" or \"real.\" A surprising number of people seemed willing to bushwhack through the Japanese knotweed and stinging nettles to find a sound buried deep in the meadow or high on top of an embankment. A challenge of sorts, Other Order could be approached as a kind of game where one tries to cover as much territory as possible, to visit each sound in its unique niche, or identify each of the plants and landmarks referenced in the sound composition and included in the project index. Related apps, including \"Alpine Garden Misguide\" by Jill Didur (2015), have successfully used the game structure of an exotic plant hunt to engage critical perspectives on the colonial histories of botanical gardens and specimen collecting. As an educational and informative piece, the authors were happy to discover that Other Order was equally appealing to adults and children, though it is less accessible to audiences with physical disabilities that would limit their movement or ability to listen through headphones. Finally, we found that visitors' appreciation of the botany of urban wilds was enhanced through the experience, but especially in the context of understanding plants in relation to their social entanglements with humans. At its core, the Other Order sound walk is an effort to combine scientific and cultural perspectives on urban ecology in a format that can reach broader publics in non-traditional settings. Bussey Brook Meadow is often mistaken for a derelict parcel of public land--a park of sorts, but with an uneasy appearance when compared to the manicured landscape of the Arboretum grounds or with sections of the Olmsted-designed Emerald Necklace of Boston parks that bear the stamp of formal landscape design. As a public artwork, Other Order is aimed at drawing visitors into the site through a sonic overlay that reveals another kind of beauty--and another perspective on ecological environments that blend the biological, technological, cultural, and social elements. The contrasting aesthetics of urban wilds and more formal parks is brought into focus, revealing a historical moment when each has undergone dramatic shifts in purpose, perception, and public use. Other Order: Sound Walk 23 TERI RUEB A map of Bussey Brook Meadow showing the locations of the fifty-five sound regions that make up Other Order. Herb Nolan, a longtime Arboretum supporter, listening to Other Order. 24 Arnoldia 75\/1 ? August 2017 NANCY ROSE Other Order is available for free download from the App Store and Google Play Store. Visitors are encouraged to download the app in advance of their visit and wear headphones in order to appreciate the stereo, binaural recordings as they blend with the actual environmental sounds of the meadow. The combination of stereo, binaural recordings with the ambient sounds that surround visitors as they move through the meadow creates an uncanny sense of being simultaneously \"here and now\" and \"there and then\" in the site, further emphasizing the complex temporalities of the meadow, and complicating the emplacement of visitors as situated actors within its operations. Urban vegetation provides autumn color along Blackwell Path in Bussey Brook Meadow. Other Order: Sound Walk 25 Acknowledgements The final work included over two and a half hours of edited sound recordings that were broken into roughly thirty-six different sound regions spread throughout the 25-acre site. Spoken elements included excerpts from conversations with the following Arnold Arboretum staff members: Ned Friedman, Maggie Redfern, Jim Papargiris, Nima Samimi, Susan Hardy Brown, Ailene Ettinger, and Bob Mayer (Arboretum volunteer). From outside the Arboretum the following people were recorded: John Lee, Eugenie Beal, Nina Brown, Lucy Hutyra, Steve Decina, Matthew Battles, Kyle Parry, Anya Yermakova, and Richard, a longtime resident of Bussey Brook Meadow. And finally, thanks to Ernst Karel for help with sound recording. Funding for the development of Other Order was provided by a generous donation from Janine Luke in memory of Melvin Seiden. References Arnold Arboretum of Harvard University. Bussey Brook Meadow. https:\/\/www.arboretum.harvard.edu\/ plants\/featuredplants\/bussey-brook-meadow\/ (accessed March 20, 2017). Boston Redevelopment Authority (BRA). 1976. Boston Urban Wilds: A Natural Area Conservation Program. Boston: Boston Redevelopment Authority. Bird, C. J. 2014. Boston's Urban Wilds: The Persistence of an Idea Over Time. Masters Thesis, City Planning, Massachusetts Institute of Technology, Cambridge Massachusetts. Burkholder, S. 2012. The new ecology of vacancy: rethinking land use in shrinking cities. Sustainability 4: 1154?1172. Carroll, S. P. 2011. Conciliation biology: the ecoevolutionary management of permanently invaded biotic systems. Evolutional Applications 4: 184?199. Daniel, T. C. et al. 2012. Contributions of cultural services to the ecosystem services agenda. Proceedings of the National Academy of Sciences 109: 8812?8817. Del Tredici, P. 2010a. Wild Urban Plants of the Northeast: A Field Guide. Ithaca, New York: Cornell University Press. Del Tredici, P. 2010b. Spontaneous urban vegetation: reflections of change in a globalized world. Nature and Culture 5: 299?315. Desimini, J. 2015. Deciphering the urban wild: remnant and re-emergent. In: A. Berrizbeitia (Ed.), Urban Landscape, pp. 163?170. London: Routledge. Didur, J. 2015. Alpine Garden Misguide, mobile app available on iTunes, June 2015. Jorgensen, A. and R. Keenan (Eds.). 2012. Urban Wildscapes. London: Routledge. Garvin, E., C. Branas, S. Keddem, J. Sellman. 2012. More than just an eyesore: local insights and solutions on vacant land and urban health. Journal of Urban Health 90: 412?426. Gulachenski, A., B. M. Ghersi, A. E. Lesen, and M. J. Blum. 2016. Abandonment, ecological assembly and public health risks in counter-urbanizing cities. Sustainability 8: 491. Keil, A., 2005. Use and perception of post-industrial urban landscapes in the Ruhr. In: I. Kowarik and S. K?rner (Eds.), Wild Urban Woodlands, pp. 117?130. Berlin: Springer. Kowarik, I. and S. K?rner (Eds.). 2005. Wild Urban Woodlands. Springer, Berlin. Hofmeister, S. 2009. Nature running wild: a socialecological perspective on wilderness. Nature and Culture 4(3): 293?315. Nassauer, J. I. and J. Raskin. 2014. Urban vacancy and land use legacies: a frontier for urban ecological research, design, and planning. Landscape and Urban Planning 125: 245?253. Pataki, D. E. et al. 2011. Coupling biogeochemical cycles in urban environments: ecosystem services, green solutions and misconceptions. Frontiers in Ecology and the Environment 9: 27?36. Pfeiffer, J. M. and R. A. Voeks. 2008. Biological invasions and biocultural diversity: linking ecological and cultural systems. Environmental Conservation 35: 281?293 Rink, D. 2005. Surrogate nature or wilderness? Social perceptions and notions of nature in an urban context. In: I. Kowarik and S. K?rner (Eds.) Wild Urban Woodlands, pp. 67?80. Berlin: Springer. Robinson, S. L., and J. T. Lundholm. 2012. Ecosystem services provided by urban spontaneous vegetation. Urban Ecosystems 15: 545?557. Rueb, T. and P. Del Tredici. 2014. Other Order: A Bussey Brook Meadow Sound Walk, Version 1.1 (275 MB). Mobile app commissioned by the Arnold Arboretum of Harvard University and available on iTunes and Google Play, October 2014. Tanner, O. 1975. Urban Wilds. New York: Time-Life, Inc. Thompson, C. W. 2012. Places to be wild in nature. In: A. Jorgensen and R. Keenan (Eds.), Urban Wildscapes, pp. 49?64. London: Routledge. Weber, F., Kowarik, I., and S?umel, I. 2014. A walk on the wild side: perceptions of roadside vegetation beyond trees. Urban Forestry and Urban Greening 13: 205?212. Peter Del Tredici is the former Director of Living Collections at the Arnold Arboretum and now a Retired Senior Research Scientist. Teri Rueb is a Professor in the Department of Media Study at the University at Buffalo -- State University of New York. "},{"has_event_date":0,"type":"arnoldia","title":"Floral Clocks, Carpet Beds, and the Ornamentation of Public Parks","article_sequence":3,"start_page":26,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25622","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14eb76b.jpg","volume":75,"issue_number":1,"year":2017,"series":null,"season":null,"authors":"Andersen, Phyllis","article_content":"Floral Clocks, Carpet Beds, and the Ornamentation of Public Parks M unicipal parks are the last territory of the decorative gardening tradition of bedding out: the practice of using brightly colored, low-growing flowering and foliage plants in ornamental patterns in beds, mounds, pyramids, floral clocks, commemorative plaques, and threedimensional figures. Arranging plants to create decorative patterns is a convention of garden design from the eighteenth century, found from the parterres de broderie of Versailles to the boxwood fleur de lis of George Washington's Mount Vernon The geometric patterns and colorful flower and foliage plants typical of bedding out garden. But in the nineteenth are seen in this postcard depicting part of Forest Park in Springfield, Massachusetts. century, perhaps as a reaction who doesn't. Bedding out of brightly colored to the long reign of the picturesque model, flowers in artificial situations became part of which favored naturalistic design, bedding out the larger discussion in which popular taste jumped the walls of the aristocratic garden and was defined as bad taste, the highbrow\/lowbrow found a home in public parks on both sides of remnants of that discussion still being argued the Atlantic. The bedding out practice elicited in gardening circles today. Bedding out was the the admiration of the public and the sustained territory of gardeners rather than landscape scorn of many landscape critics. designers, anonymous individuals whose skills A variety of names have been assigned to were admired but whose names are unknown. this practice: carpet bedding, mosaiculture, The great bedding out schemes in public parks pattern gardening, \"gardenesque.\" It is also in American cities were associated with civic called Victorian gardening, an homage to its pride, with a populist enthusiasm for both the popularity in nineteenth century Great Britain, where advances in greenhouse technology intricate floral displays and the intensive labor and the introduction of tropical and subtropical that was needed to both create and maintain species created a new way of displaying flowthem. Horticulture, as well as city beautificaering plants. Bedding out, now the shorthand tion, is inherently competitive. Supported by term, occupies a territory between art and park commissioners and local officials, municipal gardeners were encouraged to expand their craft. It is part of the history of ornamentation as well as the history of gardening. Both floral displays to accommodate the tastes of the embrace the power of serial imagery and the people--\"to show you care.\" creator's virtuosity in creating original forms. Bedding out is temporary and labor intensive--a sink-hole of energy consumption. It is Bedding out evoked heated discussion on the profoundly artificial, appealing most directly to definition of taste: what is it, who has it, and COURTESY OF THE AUTHOR Phyllis Andersen COURTESY OF UNIVERSITY OF WISCONSIN Floral Clocks, Carpet Beds 27 A color plate in Robert Thompson's The Gardener's Assistant (1878 edition) shows carpet bedding patterns complete with lists of plants to be used. NANCY ROSE 28 Arnoldia 75\/1 ? August 2017 Robinson went on to call bedding out \"pastrymaking.\" The popular and widely published British writer Shirley Hibberd called ribbon beds (long meandering beds with alternating bands of floral color) \"eels in misery.\" Landscape architect Frederick Law Olmsted's antipathy to floral display is well known. Writing in 1892 to his associates in Brookline, Massachusetts, about detached floral beds in London parks, he observed \"I have hardly seen anything yet of that kind that did not seem to me childish, vulgar, flaunting, or impertinent, out of place and discordant with good general effect.\" In an article in a 1908 issue of Ladies Home Journal, the writer blamed municipal gardeners for creating \"veritable pimples on the face of Nature.\" The Roots of Bedding Out Brightly colored flowers and foliage along with exotic tropical plants are still a common feature in city parks, such as this streetside planting in Victoria, British Columbia. the senses rather than to the power of reflection or solitary contemplation. It is not a simulacrum of nature. It is antithetical to a prevailing notion of public parks based on a pastoral model, famously invoked in Frederick Law Olmsted and Calvin Vaux's design for Central Park in New York. It did not claim to bring the country into the city. The practice of bedding out produced no theoretical treatises, no literary or painterly allusions. It tapped into the public's love of spectacle and novelty, its appreciation of skilled labor well executed. Bedding out captured the lure of the exotic by using newly discovered plants from South America and Africa, tropical and subtropical natives brought into a temperate climate. It had the repetitive power of a military parade, an analogy not lost on the British garden writer William Robinson, who observed \"Gardeners were not so much plant stewards as drill sergeants.\" Critics of this type of floral display reached new heights of rhetorical disdain. William The evolutionary process that advanced the nineteenth century version of bedding out is traced to the writings of landscape designer and writer John Claudius Loudon (1783?1843) who, in the 1830s, introduced the word \"gardenesque\" to the vocabulary of landscape. He encouraged his readers to think beyond the picturesque to what he defined as \"scientific,\" collecting plants from all over the world to test their adaptability to different climates and growing conditions (a close definition of arboreta and botanical gardens). The goal was not to imitate nature. While Loudon valued artifice and offered bedding designs in many of his publications, he did warn against the extremes of bedding out, the distorted beds and clashing colors. Loudon also recognized the limited educational opportunities for gardeners whose only option was a long apprenticeship that isolated them from new plant introductions and planting techniques. Loudon published Self Instruction for Gardeners in 1815, the first of several publications in the nineteenth century that attempted to codify best practices for both estate gardening and later municipal park management. With printing costs dropping, a number of magazines were founded that addressed professional gardeners, giving them access to information on new bedding plants and propagation techniques. They came to serve as pattern books for floral designs--fashion magazines for BIODIVERSITY HERITAGE LIBRARY MISSOURI BOTANICAL GARDEN Floral Clocks, Carpet Beds 29 With their bright colors and exotic, pouchlike flowers, South American calceolarias fit perfectly in the bedding out trend. Illustration of Calceolaria pisacomensis from Curtis's Botanical Magazine (Volume 93, plate 5677), 1867. Illustration by D. Bois of zonal pelargonium (Pelargonium zonale) and several hybrid selections (note the hybrids' larger petals and denser flower heads) in Edward Step's Favourite flowers of garden and greenhouse (Volume 1, Plate 54), 1896?1897. aspiring enthusiasts. Florists' Journal, Gardeners' Chronicle, and Gardener's Magazine were available in Great Britain, and Magazine of Horticulture, Gardener's Monthly, Genesee Farmer and Gardener's Journal informed gardeners in the United States. If principles of romanticism and aesthetic theory provided a structure for the pastoral park, advances in science and technology stimulated the expansion and complexity of bedding out. To underscore the artificiality of the bedding out system, the plant species used were often imports from South America, Africa, and the Mediterranean region. The botanical bounty collected by plant explorers, perhaps more appropriately called flower hunters, was given to botanic gardens and to commercial nurseries where species were hybridized to create showy selections with features such as compact growth, larger flowers, more brilliant colors, and variegated foliage. Plants were as much a product of the nursery trade as they were of plant collecting. Many global imports-- begonias, calceolaria, echeveria, caladiums, cannas, coleus, and more--were commonly used in bedding out configurations. Sedums, sempervivums, and other succulents also had a brief period of popularity. Palms, yuccas, crotons, monkey puzzle trees, and banana plants, all valued for their exotic forms, were brought in to serve as backdrops for theatrical staging and to punctuate the flatness of planting beds. Floral Clocks COURTESY OF THE AUTHOR The association of plants with the passing of time has a very long history. Carl Linnaeus developed an idea for a flower clock in his 1751 treatise Philosophia Botanica. Based on his field observations, he proposed a Horologium Florae, a clock using forty-six flowers which opened and closed as the day progressed. But this more literal interpretation of flowers and time evolved into decorative objects: flower plantings with an imbedded clock mechanism. In the early twentieth century the floral clock was reinvented as a decorative object for parks, tourist sites, and international expositions. COURTESY OF NIAGARA PARKS First created in 1903, the floral clock in Edinburgh, Scotland, is still a popular attraction (postcard from the early twentieth century). The design for the 40-foot-wide floral clock in Ontario's Niagara Parks is changed yearly and requires 15,000 to 20,000 bedding plants. Floral Clocks, Carpet Beds 31 Growing and Designing With Bedding Plants By the mid-nineteenth century, glass houses, once a luxury of estate gardens, became accessible to municipalities and commercial nurseries. In Great Britain, the repeal of the glass tax in 1845 dropped the cost of the material and fueled experiments with mass production. In both the United States and Great Britain advances in cast and wrought iron construction developed for the glass pavilions of the Crystal Palaces in London, Syndenham, and New York City were adapted to smaller glass structures. Magazines for gardeners offered advice on ventilation, humidity control, and heating alternatives. Commercial nurseries created acres of glass houses for the mass production of bedding plants. Nurseryman and author Peter Henderson (1822?1890) started with a small shop in New York City selling seeds. By the 1850s, his business skills and ability to predict the horticultural market allowed him to build extensive greenhouses near Jersey City, New Jersey. By his books, aimed at both the professional gardener and the amateur, and by his color catalogs, he developed a market for bedding plants, including his own introductions, most notably zinnias from Mexico and his hybrid `Giant Butterfly' pansy. Henderson visited England in 1885 and noted that the carpet style beds \"were interest- ing to the people in a way that no mixed border could ever be.\" He also noted the conspicuous lack of ornament in Central Park and Prospect Park, an omission he attributed to \"a lack of taste in the management of our public parks.\" Color theory, the investigation of human color perception, guided gardeners in the design of beds and created the distinctive intense impact of color combinations, either gaudy or brilliant according to your taste. One of the first explorations of color perception was a 1743 treatise by the French naturalist, Georges-Louis Buffon, followed by Johann Wolfgang von Goethe's Theory of Colours published in English in 1840. But it was the work of Michel Eugene Chevreul (1786?1889), a French chemist employed by the Gobelins Tapestry Works whose work on color, first directed to the textile industry but also to horticulturists, gardeners, and artists, that proved the most influential. BIODIVERSITY HERITAGE LIBRARY No plant group was more subject to manipulation than the pelargoniums (Pelargonium), which are often called (erroneously, British gardeners would say) geraniums in the United States. Native to South Africa, pelargoniums are still ubiquitous garden plants: drought resistant, blooming throughout the summer, a plant that has become a symbol of cheerful welcome in window boxes and entry planters. Zonal pelargoniums, introduced in the late eighteenth century, are characterized by alternating bands of dark and light green on their leaves and large, brilliantly colored flower heads. Continual experimentation with hybridizing various Pelargonium species resulted in hundreds of upright, prostrate, variegated, and ivy-leafed cultivars. Instantly recognizable by the general public, the pelargonium is still among the most popular bedding plants in municipal parks. The 1900 autumn catalog from Peter Henderson and Company offered tulip bulbs for bedding out patterns. BIBLIOTH?QUE NATIONALE DE FRANCE 32 Arnoldia 75\/1 ? August 2017 Michel Eugene Chevreul's color circle was used by horticulturists and garden designers when creating bedding out displays. His book, The Principles of Harmony and Contrast of Colours and Their Applications to the Arts, published in English in 1854, enhanced the gardener's understanding of how colors are modified when placed next to each other, in contrast to how the color is perceived when observed alone. Chevreul's Color Circle, a circular chart organizing complimentary colors opposite each other, was a reference guide for gardeners well into the twentieth century. Bedding Out in City Parks The city of Chicago engaged some of the best landscape architects in the country to transform the flat terrain of the city into a sophisticated park system to rival those of Eastern cities. Frederick Law Olmsted and Calvert Vaux, H. W. S. Cleveland, and later Jens Jensen, with his passionate commitment to the prairie landscape, all left their imprint on the city. But parallel to their planning work, the city encouraged ornamental planting in the form of elaborate bedding out schemes. In the 1890s Chicago built a large glass conservatory in Lincoln Park for the display of tropical plants, with extensive plant propagation areas for bedding plants. Earlier Chicago's South Park Commissioners supported ornamental plant attractions in Washington Park that included a twentyfoot-diameter globe, a sundial of echeverias, and wire structures covered in flowering plants depicting President Grant and Uncle Sam-- what one writer called \"floral masterpieces.\" In 1872 a Board of Botanical Directors was formed under the direction of H. H. Babcock, a prominent botanist and member of the Chicago Academy of Science. In a move antithetical to Olmsted, Cleveland, and Jensen's native plant perspective, the Board sent requests to botanical gardens and noted horticulturists all over the world and received seeds and bulbs from the United States, Europe, India, and Australia. Many were eventually planted out in the Chicago parks. In 1891, the journalist Charles Pullen wrote extensively on the development of Chicago's park and parkway system, especially of Olmsted's plans. He carefully threaded his way through the controversies of natural and artificial but commented that \"it is hoped that with the gradual evolution of the grander and simpler elements of the park landscape these features of curiosity will be given to more appropriate places, less antagonistic to the pleasures obtained from natural scenery.\" Boston's Public Garden, still admired for its commitment to the bedding out tradition, rests on a set of artificial conditions that eliminated any call for a rural landscape model. The Garden was created out of brackish tidal flats as part of the landfill project that created Boston's Back Bay. The supporters of the Public Garden were men with strong horticultural interests as well as a dedication to civic improvement. William Doogue, the Irish-born horticulturist hired to bring architect George Meacham's original 1859 plan for the Public Garden to life, developed flower adornments for the Garden of great public appeal. He maintained a municipal greenhouse that produced thousands of plants for the extensive beds he created throughout the CHICAGO HISTORY MUSEUM Floral Clocks, Carpet Beds 33 CHICAGO HISTORY MUSEUM A park with bedding out displays along Drexel Boulevard in Chicago. A stereo view card with images of large floral sculptures in Chicago's Washington Park. COURTESY OF THE AUTHOR Garden. A newspaper article in 1888 described the summer scene: acanthus, pyrethrum, beds of silverleaf geraniums and pansies, edged with lobelias and alternanthera. In the midst of this blaze of color, Doogue created a cactus bed, an exotic sight to Garden visitors. In one of the more memorable horticultural disputes of the nineteenth century, played out in the pages of Garden and Forest magazine in the 1880s, Mr. Doogue's plantings and their accompanying popularity with the public were condemned by both signed and unsigned Early twentieth century postcard shows bedding out in Boston Public Garden. articles in the publication. Doogue and the supporters of his distinctive floral displays were pitted against an impenetrable fortress of opposition from the likes of landscape writer and architectural critic Mariana Griswold Van Rensselaer, Arnold Arboretum director Charles Sprague Sargent, and Frederick Law Olmsted. Doogue scoffed at their limited views, their isolated lives, and, most importantly, their lack of empathy for the taste of the general public. The bedding out tradition is seen as a historical remnant of the Victorian era, outside the canon of landscape design history: at best, charming and whimsical, at worst an affront to good taste and the sanctity of a natural landscape. It is seen A Boston Public Garden floral carpet bed depicting the seal of the American as a vernacular tradition perpetuated Legion in 1930. by gardeners rather than professional evergreen hedges of about 2-foot height ... The landscape designers or landscape architects. In type of plant, height, distance apart, and planta postscript to the bedding out tradition, the ing details would be under the direction of a Philadelphia Museum of Art sponsored an art botanist and the maintenance by a gardener.\" installation in 2012 by the minimalist artist It was a short term work, installed in 2012 and Sol LeWitt (1928?2007). The work was based dismantled in 2015. It, perhaps unintentionally, on a proposal LeWitt made to the Fairmount reiterated the original power of bedding out: the Park Art Association in 1981. The resurrected appeal of geometric forms, the intervention of Lines in Four Directions in Flowers was created from LeWitt's initial instructions: \"To blocks of color in a green field, the fascination plant flowers of four different colors (white, with observable change. yellow, red, and blue) in four equal rectanguToday, we still identify public parks with lar areas, in rows of four directions (vertical, the core of civic life. The binary of ornamental horizontal, diagonal right and left) framed by versus pastoral is still rightfully argued, but, COURTESY OF THE AUTHOR 34 Arnoldia 75\/1 ? August 2017 COURTESY OF THE PHILADEPHIA MUSEUM OF ART Floral Clocks, Carpet Beds 35 Artist Sol LeWitt's work, Lines in Four Directions in Flowers, was installed in front of the Philadelphia Museum of Art from 2012 to 2015. more challenging, is the question: How do you translate planting techniques of mass appeal with contemporary values of sustainability and the still unspoken ideas of taste? In 1856, horticulturist and landscape designer Andrew Jackson Downing argued that the public park could modify artificial barriers of class, wealth, and fashion--a notion that is still valid and still contentious. Sophisticated observers may still feel a degree of discomfort at the use of bedding plants to spell out town names, patriotic emblems, comic characters--the definition of kitsch being \"the adaptation of one medium to another.\" But in the words of the art historian Tomas Kulka, \"If works of art were judged democratically--that is, according to how many people like them--kitsch would easily defeat all of its competitors.\" Further Reading Bluestone, D. 1991. Constructing Chicago. New Haven: Yale University Press. Chevreul, M. R. 1987. The Principles of Harmony and Contrast of Colors and Their Applications to the Arts. Revised edition with introduction and commentary by F. Birren. West Chester, Pennsylvania: Schiffer Publishing. Elliott, B. 1986. Victorian Gardens. London: Batsford. Musgrave, T. 2007. The Head Gardeners: Forgotten Heroes of Horticulture. London: Aurum Press. Wilkinson, A. 2007. The Passion for Pelargoniums: How They Found Their Place in the Garden. Gloucestershire: Sutton Publishing. Phyllis Andersen is a landscape historian and former director of the Institute for Cultural Landscape Studies of the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Uncommon By Any Name: Acer pensylvanicum","article_sequence":4,"start_page":36,"end_page":37,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25625","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14e816d.jpg","volume":75,"issue_number":1,"year":2017,"series":null,"season":null,"authors":"Hetman, Jon","article_content":"Uncommon By Any Name: Acer pensylvanicum Jon Hetman O ne thing I found challenging when I first started working at the Arboretum was learning (and using) scientific names for plants instead of their common names. While perhaps easier to use than those tongue-twisting Latin binomials, common names prove problematic for identification because they can refer to generic groupings (think of honeysuckle or rose) and can vary in usage from place to place (in the United States, a Tilia is called a linden; in the United Kingdom, a lime). Nevertheless, common names can offer intriguing clues about plants and their formal, natural, and historical associations. Consider the diversity of references suggested by the many common names for Acer pensylvanicum--from striped maple to whistlewood--and you begin to appreciate how one plant can inspire many appellations. Native to North America from Nova Scotia to Wisconsin and south through the Appalachians to northern Georgia, A. pensylvanicum is called striped maple or snakebark maple because its smooth, olive-green bark bears bright green and white vertical striations. It shares this trait with more than a dozen other maples in Section Macrantha, though all the rest (including A. davidii, A. maximowiczii, and A. rufinerve) originate from Asia, making the snakebark maples a great example of the eastern Asia\/eastern North America disjunct pattern of biogeography. The considerable ornamental interest provided by its bark makes A. pensylvanicum a real stand out, particularly in the winter landscape. In spring, the leaves of A. pensylvanicum unfold tinged with pink and mature to bright green. It bears large, serrately margined leaves that measure up to seven inches (18 centimeters) across. Long-stalked and typically with three sharp-tipped lobes, the leaf shape suggests a third common name for the tree, goose-foot maple. In autumn, leaves turns a clear yellow. A primarily dioecious plant with male and female flowers on different plants, the tree bears long, pendent racemes of delicate, pale yellow-green flowers in early spring, which give way to graceful chains of pinkish samaras (winged seeds) that are extremely showy by summer's end. Not overly abundant in the wild, striped maples grow to only 30 to 40 feet (9 to 12 meters) in height and spread, and are often multi-trunked because of wildlife browsing. In addition to feeding on the tree's soft shoots and young foliage, deer and moose also rub the velvet on their antlers against the smooth trunks of A. pensylvanicum as they approach the rutting season, suggesting two additional common names--moose maple and moosewood. When cultivated in the landscape with good soil, adequate moisture, and at least partial shade, moosewood can thrive as a striking specimen of intermediate size. While the species is not prone to any significant insect pest or disease problems, gardeners should protect its soft trunk from lawnmower injuries and other mishaps. The ease of its wood to yield to the knife once made it a popular choice for making whistles, and some still call it whistlewood. The Arboretum has cultivated A. pensylvanicum since 1874. Today, you may observe 14 individuals of the species representing nine accessions, including two specimens of the cultivar `Erythrocladum', selected for the coral pink to red color of its young winter twigs. Holdings of the species include individuals wild-collected by Senior Research Scientist Emeritus Peter Del Tredici in 1979 (in West Cornwall, Connecticut), Keeper of the Living Collections Michael Dosmann in 2008 (in Franklin County, New York and Mt. Wachusett, Massachusetts), and Manager of Plant Records Kyle Port in 2013 (in Orland, Maine). Most grow on the east side of Meadow Road in the Azalea Border and along the edge of the Maple Collection, and on the west side of Meadow Road in the North Woods. Visit any time of year to appreciate this truly remarkable native, no matter what name you call it. Jon Hetman is Director of External Relations and Communications at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23455","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14ea76e.jpg","title":"2017-75-1","volume":75,"issue_number":1,"year":2017,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Wardian Case: How a Simple Box Moved the Plant Kingdom","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25619","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14eb328.jpg","volume":74,"issue_number":4,"year":2017,"series":null,"season":null,"authors":"Keogh, Luke","article_content":"The Wardian Case: How a Simple Box Moved the Plant Kingdom Luke Keogh I ECONOMIC BOTANY COLLECTION, ROYAL BOTANIC GARDENS, KEW n 1873, before a single road was laid at the Arnold Arboretum, founding director Charles Sprague Sargent packed a Wardian case full of ferns from the American West. Wardian cases--wood and glass boxes shaped like small, moveable greenhouses--were used for transporting live plants. They were often more delicate than typical cargo, so to ensure protection Sargent sent the box to England with a friend who also happened to be sailing across This Wardian case dates to around 1870. the Atlantic. He hoped the ferns, from California and Colorado, might impress his colleagues at the Royal Botanic Gardens, Kew, in particular the director, Joseph Hooker. Sargent wanted to make sure that Kew's collection of North American ferns was \"as complete as possible.\" In the April 2, 1873, letter that accompanied the Wardian case Sargent wrote to Hooker: \"If ... you have all you want, will you kindly send them on to the Jardin des ARCHIVES OF THE ARNOLD ARBORETUM The Wardian Case 3 live plants around the globe had become extensive. This had only become possible because of the invention of the Wardian case. The letter also shows that botanists, arborists, agriculturists, and horticulturists from around the globe needed to remain in constant contact. Indeed, they could use one contact to begin a dialogue with another, in this case using Hooker to re-open contact with Paris. At all times these networks relied on reciprocity--one institution sends some plants and the receiver sends some back. One of the key ways a new contact was smoothed out, or an already established contact was asked for new plants, was to send them some from your own collection. Sargent concluded his letter to Hooker by sending a list of desiderata--a list of plants that they needed at Harvard. He was not only hoping for plants for the new arboretum in Jamaica Plain but also plants for the botanic garden that then existed in Cambridge. Among the most desired were palms and agaves. \"[A]s the Garden is so destitute of them that anything you can send will be most acceptable,\" wrote Sargent. A few months later Hooker was able to satisfy Sargent's request by sending palm seeds plus some very good seeds of the Abyssinian or Ethiopian banana (Musa ensete, now known as Ensete ventricosum). This was a very rare plant to have in North America at the time because it did not travel well. In the same consignment A hand-painted lithograph showing Musa ensete (now known as Hooker also included a Wardian case full Ensete ventricosum) growing in Abyssinia (a former kingdom in what of rhododendrons (Rhododendron). In the is now Ethiopia), from Curtis's Botanical Magazine, January 1861. following months Sargent would go on to Plantes at Paris.\" Sargent hoped that Kew could return the favor and send another Wardian case send at least a few plants on to Paris: \"I am sorry of ferns to Hooker. And so the reciprocal relationship continued between them. to trouble you in this way, but unfortunately we have not as yet any sure way of transporting The Challenge of Sending Plants living plants to Paris and I doubt not you are in Many common horticultural or agricultural constant communication with Dr. Decaisne.\" species, for example the Japanese umbrella pine Sargent's letter shows us a number of processes in motion during that time. While (Sciadopitys verticillata) or even tea (Camellia moving plants still relied on an important netsinensis), required the efforts of plant hunters work of botanists, by the 1870s the trade in and travelers to discover and introduce them. Just keeping plants alive on a long sea voyage was a challenge for early travelers. As early as the seventeenth century plants arrived in Europe from around the globe, to the pleasure of plant enthusiasts. As the centuries wore on, more and more rare and exotic plants steadily made their way to Europe and North America. While some plants were sent as seeds or cuttings, many plants could not be transported in those forms and had to be sent as live specimens. For naturalists this amounted to a great chalWardian cases were critical to the movement of economically important plants lenge. As shipping increased in such as tea (Camellia sinensis). A tea plantation in Tanzania is seen here. the early nineteenth century many in London it came down to the type of and the world became increasingly connected case plants were sent in. through exploration and trade, transporting live At this time naturalists and gardeners turned plants was still very difficult. their efforts to discovering the best way to In 1819, John Livingstone, the keen botanist send live plants around the globe. Was it in a and surgeon posted in Macao for the East India wooden box? Or were there other ways? Many Company, wrote to the Royal Horticultural methods for successfully moving live plants Society on the challenge of sending live plants to destinations well beyond their native range from China to London. Livingstone estimated were being tried. Accompanying Lindley's paper that only one in one thousand plants survived to the Horticultural Society were designs for the journey. He proposed a number of plans a glazed box that had been sent to him by the for the successful movement of plants. One Governor of Mauritius, Robert Farquhar. One was quite simply to send a gardener with any of the more interesting methods proposed was dispatch of live plants to make sure they were that of Nathaniel Wallich, pioneering botanist properly cared for on the voyage. But whatever and surgeon for the East India Company, based the method was, Livingstone concluded in in Calcutta; he sent a box that had a roof made his November 16 letter to the Society, that it with translucent shell inserts, which allowed \"becomes a matter of importance to attempt light in. These early decades of the nineteenth some more certain method gratifying the English horticulturist and botanist, with the plants century were an intense period of experimentation in sending live plants. of China.\" John Lindley, also of the Horticultural Society of London, described the great Ward's Glass Case challenge and care needed in sending live plants Most inventions do not come about in a vacacross oceans. In 1824, Lindley wrote, \"The uum, and so it was with the Wardian case. Many idea which seems to exist, that to tear a plant plant transportation containers, some of which from its native soil, to plant it in fresh earth, were quite successful, paved the way before to fasten it in a wooden case, and to put it on the actual Wardian case was invented. It was a board a vessel under the care of some officer, is simple case made of wood and glass and takes sufficient, is of all others the most erroneous, its name from its inventor, Nathaniel Bagshaw and has led to the most ruinous consequences.\" Ward, a London physician with a keen interest Lindley proposed a more controlled and concerned approach to sending plants. Indeed, for in the natural world. Ward's improvement on CHUCK BARGERON, UNIVERSITY OF GEORGIA, BUGWOOD.ORG 4 Arnoldia 74\/4 ? May 2017 The Wardian Case 5 Sketch of the box used by Sir Robert Farquhar to transport plants from Mauritius to London in 1824. This box holds a striking resemblance to the common Wardian case that became widely used in the nineteenth century. From John Lindley, \"Instructions for Packing Living Plants in Foreign Countries, Especially within the Tropics; and Directions for Their Treatment during the Voyage to Europe,\" Transactions of the Horticultural Society of London 5 (1824). the previous attempts was his proposal of an replanted with specimens from Australia. In airtight system in which transpiration inside Sydney the temperature was over 30?C (86?F), the case provides sufficient moisture to keep rounding Cape Horn temperatures fell to -7?C plants alive for extended periods. (We would (19.4?F), at Rio de Janeiro it reached nearly 40?C call this system a terrarium today.) (104?F), and eight months later when Mallard's In 1829, in a large sealed bottle partially filled ship travelled up the Thames the temperature with soil, Ward buried the chrysalis of a sphinx was below 4?C (39.2?F). When Ward and friend moth, with the hope that it would hatch. The George Loddiges, of the famous Loddiges & moth never flew, but he observed changes Sons nursery in Hackney, went aboard the ship inside the bottle. Sprouts of meadow grass in London they inspected the healthy fronds of a delicate coral fern (Gleichenia microphylla), (Poa annua) took life, so too did the common an Australian plant never before seen in Britain. fern Aspidium (now Dryopteris) filix-mas. Following the first successful journey to AusInstead of worrying about the moth, Ward took the sealed bottle and moved it to a window that tralia and back, Ward and his friends commenced would get the northern sun. The plants inside moving more plants in the glass cases. In 1835, survived for three years without water; in the Ward sent six cases of ornamental plants to the second year he observed the grass inside bloom head gardener for the Pasha of Egypt, and later, and the fern grew five fronds. Only after the lid following this success, coffee plants were sent. rusted and rain water entered the bottle was the George Loddiges was more ambitious. He put experiment over. into circulation over five hundred cases to all Many other experiments followed. Inside parts of the globe. It is the ingenuity and wide Ward's house was an extravagant display of use by Loddiges's nurseries that established the city gardening under glass. On March 6, 1834, Wardian case as the most compelling tool to use John Claudius Loudon, the well-known garden for transporting live plants. designer and journalist, visited Ward's house. In the nineteenth century, a Wardian case He described it as \"the most extraordinary city filled with ferns became a feature of many garden we have ever beheld.\" It was also the middle to upper class Victorian homes, includimplications of Ward's gardening in glass cases that was important. Loudon added, \"Mr. Ward has no doubt, that by boxes of this kind, with requisite modifications, he could transport plants from any one country in the world to any other country.\" At the time, Ward was in the process of testing his new glass cases on an overseas voyage. In 1833, Ward transported a perfectly packed sealed glass case containing a selection of ferns, mosses, and grasses from London to Sydney, Australia. On November 23, 1833, Ward received a letter from Charles Mallard, the ship captain responsible for the two cases: \"your experiment for the preservation of plants alive ... has fully succeeded.\" The next challenge was the return jour- Ward's case for moving plants on long sea voyages. Image from Ward's On the ney. In February 1834 the cases were Growth of Plants in Closely Glazed Cases (1852; page 71). HARVARD UNIVERSITY, GRAY HERBARIUM LIBRARY 6 Arnoldia 74\/4 ? May 2017 COURTESY OF VICTORIA AND ALBERT MUSEUM The Wardian Case 7 The Great Exhibition of the Works of Industry of All Nations was held inside the specially-built Crystal Palace in 1851. Ward's cases appeared just beside the large tree featured in the center of this illustration. ing many homes along the east coast of North America. This is where the Wardian case has largely been preserved in much historical literature--as occupying a significant place in the natural history crazes of Victorian England. By 1851 a Wardian case full of plants was exhibited at the Great Exhibition. Inside the Crystal Palace, people could view live ornamental ferns brought from far off regions; they could also view one of Ward's glass bottles with a plant in it that had apparently not been watered for 18 years. Casting our eye wider than the context of the Victorian fern craze, the scale of movement that the Wardian case facilitated is significant. Focusing on the movement of plants we see the extent to which the case was used as a unified transport technology in a time when the world was becoming increasingly connected. Ward's biggest contribution, building upon the efforts of others before him, was to propose the airtight system for keeping plants alive, as well as putting this technology into practice on many long voyages. Unlike others before him, it was also Ward's promotion of his system for transporting live plants that was important. As well as his short book On the Growth of Plants in Closely Glazed Cases (1842, republished with illustrations in 1852), he also published numerous short articles in the popular gardening media promoting his system for transporting plants. And with Ward's promotions the adoption of the case was extensive, not just by well-known nurseries like Loddiges, but by the 8 Arnoldia 74\/4 ? May 2017 British Royal Navy, French government expeditions, the Royal Horticultural Society, and many others. Travelling the Globe By 1841, Ward could add many North American ferns to his collection of global plants. He had formed a good relationship with the young Harvard botanist Asa Gray when Gray first travelled to London. Gray helped Ward accumulate an extensive collection, which he kept in glass cases in his home and put in taxonomic order using Gray's textbook. But the Wardian case wasn't just used between botanists on either side of the Atlantic. In the decades following the 1840s, botanical gardens, acclimatization societies (organizations that promoted the introduction of exotic plants and animals to see if they were adaptable), horticultural societies, and nurseries commenced wide use of the Wardian case to transfer plants around the globe. One of the most well-known uses of the Wardian case was by the Royal Horticultural Society of England. It first experimented with the cases when they sent the plant explorer Theodor Hartweg to California and central America in 1836. Following this in 1848, a member of the Society, Robert Fortune, travelled to China and successfully used Wardian cases to move tea plants from China to India. In total nearly 20,000 tea plants were transplanted in what might be one of the world's largest acts of botanical espionage. These set the foundations of the Assam and Sikkim tea industry in India. Often less known is that a This illustration of the United States Propagating Garden on the National Mall in Washington, D.C., is from the 1858 Report of the Commissioner of Patents: Agriculture. The glass houses pictured on the left and right were built to receive the tea plants that were sent to the United States in Wardian cases by Robert Fortune. BIODIVERSITY HERITAGE LIBRARY The Wardian Case 9 by the Dutch and British to be transplanted to Java and India. The rubber tree (Hevea brasiliensis) was taken from its native South America and transplanted, via Kew Gardens, to the Malay and Ceylon regions in Asia. In each of these examples the transplanted regions became leading global producers of the commodity. Many commercial crops in colonial regions were established with the help of the Wardian case. The dwarf Cavendish banana (a variety of Musa acuminata) was moved from China, via the Chatsworth Gardens in England, to the Samoan islands and spread throughout the region as a significant crop. When the French botanist Henri Lecomte was charged with establishing guttapercha (Palaquium gutta, a tree from which a useful latex was extracted) plantations in the French colonies in Indochina in the late 1800s, he took with him plants safely packed in Wardian cases. The establishment of mango (Mangifera indica) production in Queensland, Australia, also relied on the case, used as early as the late 1840s to bring grafted mango trees from India. The first Japanese plants to arrive in New England were carried in Wardian cases and delivered to horticulturists to be distributed in the Jamaica Plain (Massachusetts) area. Illustration of the gutta-percha tree (Palaquium gutta) from Franz Eugen Japanese umbrella pines, along with K?hler's 1887 publication K?hler's Medizinal-Pflanzen. dogwoods (Cornus), rhododendrons, crabapples (Malus), and cypress (Chamaecypdecade later Fortune, this time working for the United States Patent Office, sent 26,000 tea aris obtusa) survived the seventy-day journey seedlings in Wardian cases to Washington, D.C. from Yokohama to Boston, tightly and careThis instigated the United States' first experifully packed into the sealed glass cases. The first package of plants was sent in 1861 by the mental plant station in the center of the capiphysician George Rogers Hall and eventually tal, although it is suggested that not much ever found a home at Francis Parkman's small threereally happened with the tea shrubs. acre summer estate on Jamaica Pond. The folThe Wardian case was used in a range of other botanical appropriations. The cinchona tree, lowing year Hall returned from Japan with six whose bark was used in quinine-based antimamore Wardian cases. These cases were filled with many varieties of plants; among them larial drugs, was moved in secret from Bolivia NANCY ROSE 10 Arnoldia 74\/4 ? May 2017 Botanical gardens, often with a large focus on economic botany, formed hubs of distribution to move plants around the globe. The significance of Kew Gardens as a global hub of scientific knowledge and a mover of economic plants in the age of empire has been well documented in various sources. However, the significance of the Wardian case has received little specific attention. Following the invention of the case, it is estimated that in just 15 years William Hooker, director of the Gardens at Kew from 1841 to 1865, imported more plants than in the previous century. In the following eras at Kew, when his son Joseph took over as director (from 1865 to 1885), followed by William ThiseltonDyer (from 1885 to 1905), the Wardian case continued to be used extensively. From the 1860s into the twentieth century, plants were travelling to and from points including Shanghai, Ceylon, Batavia, Yokohama, Calcutta, Hong Kong, Trinidad, Tonga, Venezuela, Dominican Republic, Jamaica, Guyana, Natal, South Australia, and Melbourne. In the cases were everything from Liberian coffee to orchids, tree ferns, sisal, tonka beans (Dipteryx odorata), mangoes, and tea. The connections covered the globe and were efficient. In one letter from August 21, 1877, the collector in India, George Japanese umbrella pine (Sciadopitys verticillata) was among the first King, wrote to Thiselton-Dyer that Japanese plant species imported into New England. Seen here, Arboretum it was quicker to send plants in a accession 503-70-C grows near the Hunnewell Visitor Center. Wardian case from Calcutta to Kew than it was to get a case of plants from was Japanese honeysuckle (Lonicera japonica), Sikkim to Calcutta. which was first planted out in a nursery row in With its own imperial interests Germany was Long Island. Many plants that arrived in Wardnot to be left out of the global movement. Estiian cases have swept across North America; mates from the Berlin Botanical Gardens note some, such as the Japanese umbrella pine, have that between 1891 and 1907 over 16,000 plants become beautiful additions to landscapes and were moved by the Gardens. These included gardens, but others, like the honeysuckle, have coffee, oil palms, cocoa, rubber, and bananas. invaded woodlands across the eastern United They were moved between Berlin and VictoStates and elsewhere. COURTESY OF ROYAL BOTANIC GARDENS, KEW The Wardian Case 11 Unpacking a Wardian case at the Royal Botanic Gardens, Kew, around 1890. ria (now Limbe, Cameroon), Amani (Tanzania), Sokod? (Togo), and Simpson Harbor (today Rabaul, Papua New Guinea). German botanical gardens also played an important intermediary role when the sisal plant (Agave sisalana), cultivated for its tough fibers, was transplanted from Central America to colonies in Africa for the German East African Company. Other botanical gardens played major roles in moving plants around the globe whether it was for acclimatization, commercial, or ornamental reasons. The world renowned Jardin d'agronomie tropicale, Paris, was a major hub for acclimatization and agricultural research for the French colonies. In the first decade of the twentieth century the garden sent more than 40,000 live plants to the colonies. At Dutch gardens, including the Amsterdam Botanical Gardens and Leiden Gardens, many plants passed through on their way to Asian colonies including cinnamon, clove, mango, and ginger. For the Dutch the gardens at Buitenzorg (Bogor), Java, was the central hub for plant movement. At Russia's most prominent gardens in St. Petersburg, the German-born Carl Maximowicz became head of the botanical gardens, which COURTESY OF THE COLLECTION NATIONAAL MUSEUM VAN WERELDCULTUREN. TM-10010760 SCHLESINGER LIBRARY, RADCLIFFE INSTITUTE, HARVARD UNIVERSITY. COPYRIGHT BIBLIOTH?QUE HISTORIQUE DU CIRAD Specially crafted Wardian cases made by local Indonesian workers were used to send plants from the Buitenzorg Botanic Gardens, Java, in 1904. Workers at the Jardin d'Agronomie Tropicale [Garden of Tropical Agronomy] in Paris prepare to send live plants in Wardian cases to the French Colonies, circa 1910. With an interior botanical style inspired by the Wardian case, a rectangular terrarium sits near a New York apartment window in this photograph by Jessie Tarbox Beals, circa 1910 to 1930. The Wardian Case 13 allowed him to develop an extraordinary array of Japanese plants. In each of these places the Wardian case proved an important technology for building collections and disseminating plants to other regions. The United States became one of the most important users of the case in the early twentieth century (although there were many uses in the late nineteenth century, some of these noted above) because of the extensive usage by the United States Department of Agriculture. The Wardian case was used on many expeditions by American plant explorers importing everything from orchids to avocados and even insects. However, by the 1920s the Wardian case had become an expensive method of transport. There was also a change in the air as scientists, primarily entomologists, started to see the dangers of importing foreign plant material and biological matter. In a 1924 circular to the USDA, Beverley Galloway, the chief of the Bureau of Plant Industry, noted: \"The Wardian case, a sort of small portable greenhouse, has probably been the means of scattering more dangerous insects, nematodes, and other pests over the earth than almost any other form of carrier; hence its use is not advised except under special instructions.\" As quarantine became the order of the day, protecting the natural biota of a country became a much more significant goal of plant industries and scientists. In the eyes of gardeners, exotics still held value but they now had to compete with a greater value placed on native flora. from Australia to Boston, the case was put to good use. Collectively in the nineteenth and early twentieth century the use of Wardian cases facilitated a major plant migration across the globe. These plant distribution networks, established with the introduction of the Wardian case, are still in use today. A Century of Exchange Spongberg, Stephen A. 1990. A Reunion of Trees: the Discovery of Exotic Plants and their Introduction into North American and European Landscapes. Cambridge, Massachusetts: Harvard University Press. The Wardian case is a persuasive and widereaching example of how a simple technology for moving plants had a major impact on the ecosystems we know and value today. It was a prime mover for botanical enterprises for over a century. But by the 1940s it was largely phased out, with the last journeys carrying ornamental plants occurring in the 1960s. The case was superseded by the use of polyethylene bags and temperature controlled air transport. The Wardian case was extensively used by many different groups of people from across the globe to move many different plants; from scientists to gardeners, from ferns to bananas, Further Reading Desmond, Ray. 1986. Technical Problems in Transporting Living Plants in the Age of Sail. Canadian Horticultural History 1: 74?90. Elliot, Brent. 2004. The Royal Horticultural Society: A History, 1804?2004. Chichester: Phillimore. Galloway, Beverley T. 1924. How to Collect, Label, and Pack, Living Plant Material for Long-Distance Shipment. USDA Department Circular 323, pp. 1?12. Gardener, W. 1971. Robert Fortune and the Cultivation of Tea in the United States. Arnoldia. 31(1): 1?18. Lindley, John. 1824. Instructions for Packing Living Plants in Foreign Countries, Especially within the Tropics; and Directions for Their Treatment during the Voyage to Europe. Transactions of the Horticultural Society of London 5: 192?200. Livingstone, John. 1822. Observations on the Difficulties Which Have Existed in the Transportation of Plant from China to England, and Suggestions for Obviating Them. Transactions of the Horticultural Society of London 3: 421?29. Loudon, John C. 1834. Growing Ferns and Other Plants in Glass Cases. Gardener's Magazine. pp. 207?208. Pauly, Philip J. 2007. Fruits and Plains: The Horticultural Transformation of America. Cambridge, Massachusetts: Harvard University Press. Ward, Nathaniel B. Letters to Asa Gray, 1840?1868, Gray Correspondence Files, Archives of the Gray Herbarium, Harvard University Herbaria. (These can be read online through the Harvard Library.) Ward, Nathaniel B. 1842. On the Growth of Plants in Closely Glazed Cases. London: Paternoster Row. Luke Keogh is an environmental historian and was a Sargent Award recipient at the Arnold Arboretum in 2015?2016. "},{"has_event_date":0,"type":"arnoldia","title":"2016 Weather Summary","article_sequence":2,"start_page":14,"end_page":22,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25618","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14eaf6f.jpg","volume":74,"issue_number":4,"year":2017,"series":null,"season":null,"authors":"Pfeiffer, Sue A.","article_content":"2016 Weather Summary Sue A. Pfeiffer Winter (December 1, 2015, to February 29, 2016) This meteorological winter was the third mildest winter on record (since 1872) for Boston, yet it also brought the lowest temperatures recorded at the Arboretum in nearly 30 years and subsequent damage to a number of plants in the collections. Precipitation was above normal every month during this period for a total of 12.12 inches compared to the average of 10.40 inches. While snow typically accounts for much of winter precipitation (melted snow is measured as liquid precipitation), this year we received only 25.4 inches of snow. The snow-to-liquid ratio varies with temperature and other conditions, but assuming a rough range of 20:1 to 10:1, the snow accounted for just 1.3 to 2.5 inches of the total precipitation. This above average precipitation reduced the abnormally dry conditions of late 2015 and we entered spring under no drought conditions. An extraordinarily warm December 2015 (temperatures were 10?F above normal) led into a mild January, with only 1.5 inches of snow falling during the latter part of the month. February began with above normal temperatures--highs from the upper 50s to mid 60s--before returning to normal. Most of the snow for the month (14.7 inches total) came during two snowfall events on the 5th?6th and the 8th. During the first storm a combination of wet, heavy snow and soft, unfrozen ground led to the toppling of a number of trees, including a large 85-year-old hemlock cultivar (Tsuga canadensis `Compacta', accession 21278-A) on Hemlock Hill. By the 11th, an arctic front moved in, bringing frigid temperatures to the area. Low temperatures were below zero February 13th through 15th; the low of -11?F on Valentine's Day was the coldest ever recorded at the Arboretum for February 14th. Unseasonably warm temperatures returned for the latter half of the month. February was notable for dramatic temperature fluctuations, with a difference of 76.3 degrees between the highest (65.3?F) and lowest (-11?F) recorded temperatures. The mild temperatures throughout early winter meant buds had started to swell far earlier than usual; the mid-February freeze resulted in damaged buds on some species as well as stem dieback and complete death of some marginally hardy specimens. Spring (March 1 to May 31) Precipitation was below normal during this period, with a total of 9.52 inches compared to the average 11.57 inches. This slight decrease in precipitation lead to abnormally dry conditions as we entered summer. March saw temperature fluctuations fairly typical for spring weather. A record high was set on March 9th when the temperature soared to 78?F, breaking the old record by 6?F. This made an 89 degree temperature range within 24 days from the -11?F record on February 14th. Overall, March was warmer than average and had less precipitation than normal (2.96 inches versus 4.33 inches) despite consistent and regular rain events throughout the month. On March 31st a storm brought wind gusts in the high 30s mph. KYLE PORT An arctic system caused erratic temperature fluctuations in early April (1st to the 8th) along with 5.5 inches of snow and windy conditions. Several nights with below freezing temperatures, including a low of 18.5?F on April 4th, resulted in damage to emerging foliage, buds, and open flowers. Spring flowering ended for many magnolias, whose tepals turned to brown mush, as well as forsythias and some spring bulbs. These conditions resulted in reduced flowering on other early blooming trees and shrubs and slowed foliage expansion. Significant temperature fluctuations occurred in April and the KYLE PORT Cornelian cherry (Cornus mas) and forsythia were in bloom on March 30th, a few days before a cold front brought sub-freezing temperatures that damaged many flowers and buds. A newly accessioned Carolina jessamine (Gelsemium sempervirens `Margarita', accession 520-2016-A) was in bloom on May 3. This cultivar, `Margarita', is reported to be somewhat more cold hardy than the species typically is. KYLE PORT Visitors enjoying a beautiful day at the Arboretum on May 20th passed by a blooming weigela (Weigela praecox `Gracieux', accession 164-2000-B). monthly high of 80.5?F was recorded on April 22nd, though the month overall was cooler than average. Precipitation was normal, with most falling during the first half of the month. May began with unseasonably cool temperatures and rain each day of the first week. These conditions further delayed plant development but did extend bloom time for early spring plants and late flowering bulbs. Unfortunately the flower bud damage from the extreme cold in February and the early April freeze resulted in a less colorful spring than usual. Lilac Sunday on May 8th started cloudy and dreary with light showers, though the afternoon was sunny and warmed up into the 60s. Lilacs were also delayed in their flowering and did not peak until the following week when temperatures warmed and sunny conditions returned. Temperatures heated up during the last week of the month, hitting 95?F on the 28th, Memorial Day, which set a new record and helped to advance plant growth. A storm at the end of the month soaked the ground and brought the strongest winds of the year; fortunately there was minimal damage in the collection. Summer (June 1 to August 31) Summer 2016 was very hot and very dry. Precipitation was well below normal, only 4.05 inches compared to the average 10.44 inches, and led to moderate, severe, and eventually extreme drought conditions as we entered autumn. June was warmer than usual and precipitation was far below normal. By midJune, the effect of winter damage was evident and it became clear which plants would not leaf out and stood dead in the landscape. Unirrigated turf began to go dormant, making the landscape look more like August than June, and supplemental watering started in the collections. Cloudy conditions prevailed on only three days in June and solar radiation for the month was very high. 2016 Weather 17 KYLE PORT July was much warmer than usual; we had 14 days with high temperatures in the 90s, including an eight-day heat wave from July 21st to 28th. The hottest temperature of the year, 97.5?F, was recorded on the 25th. Precipitation was far below normal for the month. A fast moving thunderstorm on July 18 brought strong winds, storm damage, and washouts. We received almost half an inch (0.47 inches) of rain that day, but most of this was lost as runoff because the entire amount fell within a 15-minute period. Signs of drought stress became evident throughout the landscape and irrigation continued. August did not offer much reprieve. After a few pleasant days in the mid 70s, the summer heat returned with high temperatures in the 80s and 90s for the remainder of the month. On August 16, this high heat and lack of precipitation upgraded the drought from severe to extreme for the first time here since national drought monitoring began in 2000 (US Drought Monitor ? National Drought Mitigation Center). On the 22nd, an overnight front brought 0.76 inch of rain, more than half the monthly total. In the end, this was the warmest August ever recorded in Boston, with below average temperature on only four days. By the end of summer the compounding effects of defoliating insects (winter moth, gypsy moth, etc.), drought conditions, and unusually warm weather left many plants showing signs of extreme stress (green leaf drop, premature fall color, wilting, browning along leaf margins). Irrigation continued but could not make up for the lack of rainfall. Conditions had progressed to extreme drought by August 17 when Arboretum Plant Records Manager Kyle Port documented the dry slope of Peters Hill, looking toward Hemlock Hill and the Boston skyline. 18 Arnoldia 74\/4 ? May 2017 How Hot and Dry Was It in 2016? Number of days with temperatures in the 90s: 24 May: 1 day June: 1 day July: 14 days August: 7 days September: 1 day The average number of days per year with temperatures in the 90s is 13 (for Boston). The last year with more than 20 days in the 90s was 2010 (25 days). Summer 2016 (June 1 to August 31) was the second driest on record at the Arnold Arboretum. We received 4.05 inches of rain; the record is 3.97 inches set in 1957. (The official Boston weather station received only 3.92 inches, which did beat the record, making 2016 the driest summer ever recorded for the City.) PRECIPITATION SURPLUS\/DEFICIT BY SEASON Winter 2015?2016 12.12 inches average is 10.40 inches surplus of 1.72 inches Spring 2016 9.52 inches average is 11.57 inches deficit of 2.05 inches Summer 2016 4.05 inches average is 10.44 inches deficit of 6.39 inches Autumn 2016 9.92 inches average is 11.35 inches deficit of 1.43 inches 2016 CUMULATIVE PRECIPITATION DEFICIT 4 WATER (inches) 2 0 -2 -4 -6 -8 -10 WINTER SPRING SUMMER FALL 2016 Weather 19 2016 Drought Conditions at the Arnold Arboretum FROM THE UNITED STATES DROUGHT MONITOR, THE NATIONAL DROUGHT MITIGATION CENTER Scale D0............. (Abnormally Dry) D1............. (Moderate Drought) D2............. (Severe Drought) D3............. (Extreme Drought) D4............. (Exceptional Drought) Arnold Arboretum conditions (dates of status change) January 1, 2016............ D0 (Abnormally Dry) March 1, 2016............ Normal conditions June 7, 2016............ D0 (Abnormally Dry) June 14, 2016............ D1 (Moderate Drought) July 26, 2016............ D2 (Severe Drought) August 16, 2016............ D3 (Extreme Drought) November 1, 2016............ D2 (Severe Drought) In 2016, we had 14 weeks with normal conditions (no drought) and 38 weeks with drought conditions 14 weeks............ Normal Conditions 10 weeks............ D0 (Abnormally Dry) 6 weeks............ D1 (Moderate Drought) 11 weeks............ D2 (Severe Drought) 11 weeks............ D3 (Extreme Drought) In 2015, we had 28 weeks with normal conditions (no drought) and 24 weeks with drought conditions 28 weeks............ Normal Conditions 21 weeks............ D0 (Abnormally Dry) 3 weeks............ D1 (Moderate Drought) Previous years reaching Severe Drought Conditions (records began in 2000) 2012-- 2 weeks............ D2 (Severe Drought) 2002-- 5 weeks............ D2 (Severe Drought) From 2000 through 2015 there were no instances of Extreme Drought at the Arnold Arboretum. 20 Arnoldia 74\/4 ? May 2017 Autumn (September 1 to November 30) IMAGE COURTESY OF NASA Rainfall was again below average for this period, with 9.92 inches compared to the average 11.35 inches, and we entered autumn in extreme drought conditions. September's total was far below average; some relief arrived in October when a total of 5.81 inches of rain fell, which was more than the combined rainfall from June through September (5.48 inches). September was another warmer than usual month with minimal precipitation. The remains of Hurricane Hermine offered hope to alleviate drought conditions, but ultimately produced less than half an inch of rain as it passed by on the 8th. Temperatures warmed to above normal for most of the rest of the month before finally cooling down to the 50s at the end of the month. October was a very wet month, bringing soaking rains on five occasions. Torrential downpours occurred on the evening of the 21st, bringing half an inch of rain within a 15-minute period and an inch of rain over a one-hour period. This left low areas flooded and there was significant erosion on gravel paths and washouts in mulched beds. Three storms in October each delivered between 1.25 and 2 inches of rain. Despite this much-needed precipitation, extreme drought conditions persisted and much of the rainfall was lost as runoff. October temperatures were normal with pleasant fall days and cooler nights. Sourwood (Oxydendrum arboreum), ginkgo (Ginkgo biloba), red maple (Acer rubrum), sugar maple (A. saccharum), and hickories (Carya spp.) produced a great show of fall foliage color, but the leaves of many other trees desiccated or dropped off prior to color change because of the drought. Anticipated rain from a dwindling Hurricane Hermine didn't materialize. This satellite image shows a swirl of clouds with no rainfall off the coast of southeastern Massachusetts on September 8th. Arnold Arboretum Weather Station Data ? 2016 Avg.Avg.Avg.Max. Min. Precipi- SnowMax. Min. Temp.Temp.Temp.tation fall (?F)(?F)(?F)(?F)(?F) (inches) (inches) JAN 30.422.930.759.4 6.7 3.38 8.8 FEB 44.323.533.965.3-11.0 4.2415.1 MAR 50.833.642.278.118.9 2.96 2.6 APR 56.137.146.680.518.5 3.62 5.3 MAY 67.848.958.394.939.7 2.94 JUN 78.757.167.989.849.5 1.22 JUL 85.664.074.897.555.3 1.34 AUG 86.764.375.597.052.4 1.49 SEP 75.657.866.793.538.8 1.43 OCT 63.244.553.879.827.8 5.81 NOV 54.036.445.270.228.1 2.69 DEC 41.125.733.457.4 3.1 2.92 6.2 Average Maximum Temperature . . . . . . . . . . . 61.9?F Average Minimum Temperature . . . . . . . . . . . 43.0?F Average Temperature . . . . . . . . . . . . . . . . . . . . . 52.4?F Total Precipitation . . . . . . . . . . . . . . . . . . . . . . . 34.04 inches Total Snowfall in 2016 . . . . . . . . . . . . . . . . . . . 38.0 inches Snowfall During Winter 2015?2016 . . . . . . . . . 33.3 inches Warmest Temperature . . . . . . . . . . . . . . . . . . . . 97.5?F on July 25 Coldest Temperature . . . . . . . . . . . . . . . . . . . . -11.0?F on February 14 Strongest Wind Gust . . . . . . . . . . . . . . . . . . . . . 38.2 mph on March 31 Last Frost Date . . . . . . . . . . . . . . . . . . . . . . . . . . 32.0?F on April 27 First Frost Date . . . . . . . . . . . . . . . . . . . . . . . . . 27.8?F on October 27 Growing Season . . . . . . . . . . . . . . . . . . . . . . . . 183 days Growing Degree Days . . . . . . . . . . . . . . . . . . 3165 days Number of Days at 90?F or above . . . . . . . . . . . 24 days WILLIAM (NED) FRIEDMAN 22 Arnoldia 74\/4 ? May 2017 Despite the drought, some trees in the collections still displayed spectacular fall color including this black oak (Quercus velutina, accession 127-2016-A) photographed on November 19th. November was slightly warmer than normal with below average precipitation. On November 1st, drought conditions were downgraded from extreme to severe but supplemental irrigation continued to be necessary. We received regular precipitation throughout the month with fewer intense storms and a light dusting of snow on the 21st. The first half of November was unseasonably mild but more normal conditions returned mid-month and cooled further during the last week. Red oak (Quercus rubra) foliage was a beautiful red despite the ongoing drought conditions. By the end of autumn, we were 8.15 inches below normal rainfall for the year. Early Winter 2016?2017 December brought the beginning of meteorological winter. We finished the year (January 1 to December 31) with a deficit of 8.6 inches of precipitation and entered 2017 still in severe drought conditions. 2016 Recap It was a tough year for the Arboretum's living collections. Cold temperature events in February and April led to much damage and even the death of some plants. Summer brought hot and humid conditions and little precipitation, leaving soils extremely dry. Autumn saw some relief with cooler temperatures and more rain than summer, but drought conditions still prevailed. We are concerned about the long term effects of drought on the health of our collection and expect to see the effects of the 2016 water deficit on plant survival and growth in 2017. Sue A. Pfeiffer is an Arboretum Horticulturist at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Witness Tree: What a Single, 100-Year-Old Oak Tells Us About Climate Change","article_sequence":3,"start_page":23,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25621","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14eb726.jpg","volume":74,"issue_number":4,"year":2017,"series":null,"season":null,"authors":"Mapes, Lynda V.","article_content":"Witness Tree: What a Single, 100-Year-Old Oak Tells Us About Climate Change Lynda V. Mapes Environmental reporter Lynda V. Mapes spent a year at Harvard University's Harvard Forest in Petersham, Massachusetts. There, she got up close and personal with a special red oak (Quercus rubra) that provided great insights on forest life and the growing effects of climate change on the natural world. This article is adapted from her recently published book, Witness Tree: Seasons of Change with a CenturyOld Oak, which chronicles her experience. 2017. BLOOMSBURY ISBN 978-1-63286-253-2 I first met the oak in the fall of 2013, walking the Harvard Forest with John O'Keefe. A biologist given to wearing the same two sweaters all winter--that's a long time in Massachusetts--and a slouchy rag wool hat, John has walked the same circuit of 50 trees in the Forest for more than 25 years. John likes to say he started his long term survey of the timing of the seasons in the Forest, revealed in the budding, leaf out, leaf color, and leaf drop on the trees, as a way to get outside at least one day every week, then just never stopped. By now, he has compiled a valuable and unique record. Seasonal changes in nature are among the most readily observable clues to the biological effects of our changing climate, as warming temperatures reset the seasonal clock. In forests, water use, the growth rate of trees, the length of the growing season, and temperature all are connected. So John's work, documenting the seasonal gyre of the woods, was a look, told through the language of leaves, at our changing world. His foot survey is literally the ground truth for images of the tree canopy that are beamed over the Internet, continually recorded in daylight hours by surveillance cameras, watching these trees' every move, from 120 feet overhead. With John's tree-by-tree observations, the forest-level view from the cameras and other devices on observation towers, and even a drone used to fly regular photographic missions, these must be among the most closely-monitored trees in the world. For while the Harvard Forest is a natural wood, reminders that it is also an outdoor laboratory and classroom are never long out of sight. Spread over nearly 4,000 acres, the Harvard Forest, founded in 1907 and with more than 100 years of research in the archives, has one of the longest records of some types of data anywhere. Trees bristle with tags and flagging, and the Forest floor is studded with equipment. There are light sensors, and laundry baskets gathering leaf litter. Often, amid the birdsong, came sounds of science, from the buzz of a drone 24 Arnoldia 74\/4 ? May 2017 ALL PHOTOGRAPHS BY THE AUTHOR UNLESS OTHERWISE INDICATED flying a photographic mission overhead, to the hum of motors, and fans. The reality is this forest is under a microscope. It's the fulltime, year-round focus of a staff of about 40 to 45 biologists, modelers, GIS specialists, historians, ecologists, dendrologists, paleoecologists, information and communication specialists, policy experts, atmospheric chemists, research assistants, lab technicians, and administrative staff at the Harvard Forest with an operating budget from $4.5 to $6 million a year, and a larger cadre of visiting researchers from around the world. On his weekly survey walks, John measured little but the occasional snow depth or length of an unfurling leaf. But what he does do is look closely at a set number of tracked trees, chosen to represent a range of species, heights in the canopy, and forest environments--dry, wet, open, and shaded. He makes systematic notes of his observations on data sheets he created for the purpose, filled out the same way each week, year by year. Professor Andrew Richardson of Harvard University was among the first of many researchers to use John's records in influential scientific papers about the effects of climate change on forests. I met Andrew in August 2013, when I first arrived in Cambridge as a Knight Fellow in Science Journalism at MIT. I wanted to explore A spectacular luna moth rests on a building in early June. new ways to tell the story of our changing climate--a yawner of a story for too many, if told as a distant debate about treaties, dueling politics, and doomsday scenarios. The stakes are high: species extinction, the function of natural processes, and viability of habitats. But the facts won't matter if we can't get anyone to pay attention. I wanted to tell the story through the charisma of living things, and the compelling but largely overlooked drama of the delicate seasonal timing of the natural world and how it was being disrupted. So when Andrew took me up on my request to sit in with his lab, and John let me join his walks, I decided to dive deep. \"John,\" I wrote in an email not long after our first survey walk together, \"I need a tree.\" We picked it not long after--a single, glorious, nearly 100-year-old red oak in his survey that I could use as a narrative frame for my own inquiry into the Richardson lab's work. What where they learning? Could I see climate change at work in this forest, and even in this one tree? Just as settlers used notable trees, known as witness trees, to mark the metes and bounds of changing landscapes, could the big oak reveal the changing climate? John's walks were enthralling. He noticed everything, and with all five senses, creating in his field notes a portrait of the forest in Pointillist detail: how firm the tree's buds were, or whether they had softened and were getting ready to crack open. The sound of the first call of wood frogs, the scent of mineral soil as the frost melted from the ground. The sight of the leaves' first emergence; the filling and draining of puddles; the flow of the streams, and first unfolding of woodland flowers. The autumn colors of the leaves, the thunk of falling acorns; frost flowers and ice on the puddles, and the wintergreen taste of birch bark. Here was a place richly and closely observed, right down to the mud and black flies. With nothing more than a pair of binoculars, six-inch ruler, and clipboard, John, by walking the Forest again and again, amassed a detailed calendar of the seasonal year, his tiny handwriting in Number 2.5 pencil recording local events with planetary implications. His findings over the decades were clear. On average, spring is coming earlier. Fall is coming later. And winter is being squeezed on both ends. Everything in the woods reflected these changes, from the level of water in the vernal pools and springs to when the black flies were biting, the ground frozen, or leaves budding out or finally coming off the trees. It wasn't a matter of conjecture or political argument; the discussions of who does and doesn't \"believe\" in climate change in editorial pages, news reports, and Congressional debates frames this all wrong. The changing climate, trees, streams, puddles, birds, bugs, and frogs attest, is not a matter of opinion or belief. It is an observable fact. Leaves don't lie; frost isn't running for office, frogs don't fundraise, pollinators don't put out press releases. What John compiles, while taking all these walks, is the testimony of an unimpeachable witness: the natural world. Studying Phenology Discerning the workings of the natural world in seasonal timing has a long history. The roots of the word are the Greek words phaino, meaning to show or appear, and logos, to study. It's from phaino, too, that we get phenomenon, and traditionally phenology has consisted of the study of the timing of biological phenomena in nature and the relationship of these phenomena with Earth's environment, particularly the climate. The Belgian botanist Charles Morren argued that like meteorology, botany, zoology, physiology, and anthropology, this merited being a scientific discipline unto itself: phenology. He is credited with the first use of the term at a public lecture at the Belgian Royal Academy of Sciences at Brussels in 1849. Biologist John O'Keefe has recorded phenological details about trees at Harvard Forest for decades. Mushrooms, moss, and fallen leaves color the forest floor. 26 Arnoldia 74\/4 ? May 2017 Newly expanded foliage provides a bright green crown for the witness tree. MELISSA LEVANGIE Witness Tree 27 swallows and cuckoos, rooks building nests, and all manner of plant activity, from flowering snowdrops, wood anemones, and hawthorns to leafing birches, elms, oaks, beech, and horse chestnut. The recording duty passed from one generation of Marsham's descendants to another until the death of Mary Marsham in 1958. Mainstream science left phenology aside long ago. But it's being rediscovered, as researchers look for evidence of climate change in the seasonal calendar of living things. Old photographs, records of birding and garden clubs, even art and literature reveal changes subtle in the moment but visible over time. The daffodil of Shakespeare has advanced its bloom time so drastically as to no longer fit its literary frame: \"Daffodils, That come before the swallow dares, and take The winds of March with beauty,\" Shakespeare wrote in The Winter's Tale. March. Not in January. And certainly not at Christmas, as happened in 2015 when the United Kingdom witnessed its warmest start to December in 50 years, The Guardian reported. At this rate, Britain's native daffodil, the Lent lily [Narcissus pseudonarcissus]--named Author Lynda V. Mapes takes notes aloft in the century-old red oak. for its expected February?March bloom Phenology's roots are in old-style, hands-on time--is going to need a new name. Of course observation like John's, practiced long before this just confirms what the gardeners, the hikers, the outdoorsmen and women of every the term phenology was invented. The longest sort already know from their own sense of a continuous written phenological record is probably marking the first flowering of cherry trees fraying natural order. Reliable patterns of at the royal court of Kyoto, Japan, dating back nature's pageant are slipping their chronology. to AD 705. In Europe, French records of grape Phenology Plus Technology harvest dates in Burgundy stretch back to 1370, For Andrew Richardson, John O'Keefe's records and have been used by scientists to reconstruct offer valuable data he uses to explore the effects spring-summer temperatures back into the of climate change on tree physiology and Middle Ages. seasonal timing of the forest canopy. The object In England, Robert Marsham in 1736 began is to probe the forest at a variety of scales, from recording what he called his \"Twenty One Indications of Spring\" at his country estate in individual trees to the forest, region, and biosphere. The data from John's weekly walks Norfolk. He tracked the seasonal stirrings of has also helped Andrew deploy phenology as a animal life: croaking frogs and toads, singing lens on the workings of the forest in a whole nightjars, pigeons and nightingales, arriving 28 Arnoldia 74\/4 ? May 2017 new way--and brought new relevance to John's work. It all got started when Andrew was at the University of New Hampshire, working with his colleagues making measurements of the daily and seasonal rhythms of carbon dioxide exchange between the trees and the atmosphere-- the breathing of the forest. He was using instruments at the top of a 90-foot-tall tower in the Bartlett Experimental Forest in the White Mountains of New Hampshire. Then he had a hunch there were a lot of other things he could also be measuring to get a better idea of how the ecosystem Professor Andrew Richardson examines a core extracted from a tree worked. Which, in a project in Harvard Forest. meeting one day, led to a conversation with one of Andrew's collaborators. Voila: spring, pinpointed from the pixel mix. What, they wondered, about putting a camera Now the team could track the development of on the tower, with the thought that at the very the canopy all the way into summer, with every least they would get cool pictures of the forest day's incremental growth in the leaves showing canopy through the seasons for presentations up as increasing numbers of green pixels. And at science talks? come fall, the camera's pixilated signals of leaf They figured they would also probably be able coloring and drop were just as clear. Suddenly, to tell when the leaves came out and when they big swaths of landscape could be remotely monitored for seasonal development, over the Web, fell off, which would also be useful for estimating growing season length, key information for from anywhere. scientists studying how much carbon forests It was a breakthrough. Here was the possibility of creating a whole new kind of observatory: pack away. Within a few weeks they installed a remote, digital observatory, with a network what was then a state of the art camera, beaming its images over a wireless connection back of cameras that could monitor the rhythm of to a server on campus. When the first images the seasons as they transformed the land, over came in over the Internet to their computers, as large an area as the cameras could be placed, they were delighted that, dinky as it was, the with the information streamed to a central camera was performing just as they hoped. Sudserver where the data could be shared, archived, denly, they could monitor their remote field site and analyzed. Andrew dubbed it the PhenoCam from their desks. That got Andrew thinking. network. There had never been anything like it. The next summer, Andrew asked a PhD stuIn less than a year, Andrew found funding dent, Julian Jenkins, whether he thought he to start a small PhenoCam network to observe could use computer analysis to spot the beginforest phenology across northern New Engning of spring green-up in the images. In just land and adjacent Canada. That was in 2007. days, Julian created a computer program that Then the National Science Foundation (NSF) converted the red, blue, and green pixels in the in 2011 provided money that allowed the team camera image to numeric values. He then could to expand the monitoring network. Next, in count the amount of greenness in an image. 2013, NSF kicked in more money that the team Witness Tree 29 Observation towers in Harvard Forest hold cameras and other devices for research studies. used to involve volunteers in interpreting and analyzing more than 5 million images streaming into a network by then grown to some 250 sites across North America, uploading images at least once an hour, seven days a week, during daylight hours. The cameras were all over the place, from instrument towers such as those in the Harvard Forest to weather stations and building tops, from forests to tundra to Hawaiian grasslands and the desert southwest. The PhenoCam network brought the phenological tradition of Robert Marsham, Thomas Jefferson, Henry David Thoreau, and Aldo Leopold into the digital age. What would Jefferson have given for a PhenoCam on his beloved gardens, instead of having to wait for letters from Monticello to fill him in on what was in leaf and in flower. We even put a camera for the network under my red oak. Visit it at http:\/\/harvardforest.fas.harvard. edu\/webcams\/witness-tree Here was the ability to see the forest not only up close, from tree to tree, as John does, but at scale. The proverbial forest for the trees. Researchers are no longer limited only to what can be seen on foot, or the occasional imagery of a satellite, available only intermittently and from a great distance. Not surprisingly, Andrew and his collaborators are still figuring out what to do with so young a method. Their work keeps turning up surprises. New Insights on Climate Change Trevor Keenan, now at the Lawrence Berkeley National Lab, with Andrew published a paper in 2015 showing that the timing of spring and fall are connected, but not in the way widely 30 Arnoldia 74\/4 ? May 2017 The witness tree red oak stands leafless in late autumn. supposed. Conventional wisdom--and many climate models--held that the warmer temperatures that brought on an earlier spring would also mean a later fall, and a longer and longer growing season. But Trevor and Andrew found out that the timing of autumn correlates more closely with the onset of spring than with temperature or day length. Spring, it turned out, exerted a strong control on the timing of fall, somewhat offsetting the effect of warming. The findings do not imply a growing season of fixed length, as the relationship between spring onset and autumn senescence they discovered was not one to one. Rather their results suggested that current models don't include the effects of spring on autumn, leading to an over-prediction of the extension of the growing seasons by as much as 50 percent under future warming scenarios. \"It was a eureka moment,\" Trevor said. Struck by the importance of their initial findings, Trevor scaled up to investigate seasonal trends on the entire east coast. The same pattern still held true. There are several possible explanations. \"Plants know from the history of their ancestors how long their timeline is,\" Trevor said. \"So it makes sense they would have some mechanism built into their optimum function, to have a pre-programmed senescence ... The question is how quickly can they learn to change and Witness Tree 31 detect that the environment around them has changed?\" Another theory is that once trees have filled up their carbon stores they are finished with their work for the year, even though the weather is still fine. \"They have been as productive as they need to be for the year,\" Trevor said. \"They are done.\" For me the idea of seasons lasting longer than the leaves could stay on the trees was a lot to take in. There is something unnatural about it--because of course, it is unnatural. It's a human-caused forcing of the climate system, imposed on a natural physiological cycle with its own timing. There are two seasons now: the seasons of living things, and the seasons made by us. Trevor expected that in time the trees would catch up, using their ability to adapt to take advantage of longer growing seasons, as trees do further south. The question is how fast. Long term carbon sequestration measurements at the Harvard Forest also show that trees at the Forest, dominated by red oak, have been growing faster since the 1990s, as global average temperatures and carbon dioxide levels began their most rapid rise. By now, red oak is putting on more mass than any other tree species in the Forest, and faster. True, that is partly just red oak's nature. The relatively young age of the forest, still recovering from the deforestation of the nineteenth century, also makes for this strong growth. But the red oak's surge is also the result of climate change, manifest in warmer temperatures on average in winter, increased rainfall, and growing seasons lasting longer than at any point in the last two decades. With the millions of microscopic openings on their leaves, called stomata (from the Greek stoma, for mouth), trees also are speaking truth about the effects of the changing atmosphere. Water vapor, carbon dioxide, and oxygen all move in and out of leaves through these openings, creating a survival challenge. But Andrew and Trevor documented in another widely-read published paper that at higher carbon dioxide levels trees, including red oak at the Harvard Forest, are working more efficiently. They don't open their stomata as much or as often to take in the carbon dioxide they need. That means they can make as much and even more food while using less water. It also suggests a shift in the physiology of trees, with profound implications for everything from water cycling to climate. Trees like my big oak are changing their inner workings, using less water even as they put on more growth as temperatures warm and carbon dioxide levels rise. From the sky and its atmosphere to the seasonal timing and growth rate of trees and, deeper still, all the way into the photosynthetic process of individual leaves, human fingerprints are now on the most grand to the most intimate scales of our planet. You could see all this even within one tree. The big oak's witness was clear: Our world is already changing. Acknowledgements The author is grateful to the Knight Program in Science Journalism at MIT for the purchase of the PhenoCam under the witness tree at the Harvard Forest and to the Harvard Forest for keeping it online. To learn more about the witness tree project see http:\/\/harvardforest.fas. harvard.edu\/witness-tree Sources Keenan, T. F., G. Bohrer, D. Dragoni, J. W. Munger, H. P. Schmid, and A. D. Richardson. 2013. Increasing forest water use efficiency as atmospheric carbon dioxide concentrations rise. Nature 499: 324?327. Keenan, T. F. and A. D. Richardson. 2015. The timing of autumn senescence is affected by the timing of spring phenology: implications for predictive models. Global Change Biology 21: 2634?2641. Morton, O. Eating the Sun: How Plants Power the Planet. 2008. New York: Harper Perennial. Richardson, A. D., T. A. Black, P. Ciais, N. Delbart, M. A. Friedl, N. Gobron, D. Y. Hollinger, et al. 2010. Influence of spring and autumn phenological transitions on forest ecosystem productivity. Philosophical Transactions of the Royal Society, Series B 365: 3227?3246. Lynda V. Mapes is the environment reporter at the Seattle Times and the author of Witness Tree. Her research was supported by a Knight Fellowship in Science Journalism at MIT and a Bullard Fellowship in Forest Research at the Harvard Forest. For more on Lynda's experience and the book visit http:\/\/lyndavmapes.com. "},{"has_event_date":0,"type":"arnoldia","title":"Through the Seasons with Sassafras","article_sequence":4,"start_page":32,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25620","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14eb36d.jpg","volume":74,"issue_number":4,"year":2017,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"Through the Seasons with Sassafras Nancy Rose S assafras (Sassafras albidum) is an admirable tree any time of the year. Native to most of the eastern United States as well as far southern Ontario, sassafras is a mediumsized (typically 30 to 60 feet [9 to 18 meters] tall) deciduous tree with an attractive tiered branching habit. It may form dense, shrubby thickets as suckers arise from its shallow, wide-spreading, lateral root system, especially in sites like old farm fields where it has room to spread. Sassafras is primarily dioecious, bearing staminate (male) and pistillate (female) flowers on separate plants. Blooming in early to mid spring, the fragrant yellow flowers are borne in clusters that, en masse, put on quite a show despite the relatively small size (about 1\/3 inch [8 millimeters] in diameter) of individual flowers. Sassafras fruits are rather striking: ovoid, deep blue drupes cupped in fleshy, cherry red pedicels that often persist after the fruit has dropped or been eaten. Sassafras albidum has unusual foliage--its leaves may display three distinct morphologies, all of which may be present on the same tree. The three leaf shapes are 1) an unlobed oval, 2) a two-lobed \"mitten,\" with one large lobe and a smaller \"thumb\" lobe, and 3) a threelobed, trident-like form. Sassafras foliage is a pleasant-enough light green in summer, but its autumn coloration in shades of yellow, orange, and red is truly spectacular. The foliage also has a culinary aspect; fil? powder, a flavoring and thickening agent used in Creole gumbo, is made of young sassafras leaves, dried and finely ground. Sassafras is also a food plant for caterpillars of spicebush swallowtail and tiger swallowtail butterflies. Young sassafras twigs are olive green and, when scratched, emit a lemony, slightly medicinal odor. Mature bark on trunks is orangish brown and deeply furrowed. The yellowish wood is light and somewhat brittle, however, it is fairly rot resistant and so has been used to make barrels, fence posts, and other items. Oil of sassafras is extracted primarily from the bark of sassafras roots and has been used for medicines, fragrances, and flavorings, including for root beer. Sassafras is a member of the laurel family (Lauraceae), a group of mostly tropical trees and shrubs. There are just three species in the genus Sassafras. In addition to the North American S. albidum there are two similar looking species in Asia, S. tzumu from China and S. randaiense, endemic to Taiwan. Another familiar temperate region genus in the family is Lindera, the spicebushes. Scratch the stem of the native North American spicebush (Lindera benzoin) and you'll smell a lemony-medicinal fragrance similar to sassafras. For all its ornamental attributes you'd think sassafras would be more widely planted. Perhaps the main reason it's not on every street corner is that it's a bit difficult to propagate and transplant. Stem cuttings do not root readily, so propagation is done from seeds, which require stratification, or from root cuttings. Fortunately, container-grown sassafras can be found at some nurseries, especially those specializing in native plants. Sassafras grows best in moist, well-drained sandy loam but also tolerates other soils as long as they're well drained. It is generally cold hardy through USDA Plant Hardiness Zone 5 (average annual minimum temperature -10 to -20?F [-23.3 to -28.9?C]). There are currently ten specimens of Sassafras albidum growing in the Arboretum. The most prominent of these is a group (22915-A, B, C, and E) growing right along Bussey Hill Road across from the lilac collection. This is a a great-looking grove of sassafras, but unfortunately their exact provenance is unknown (they were accessioned in 1950 as \"existing plants\"). We do have several accessions of known provenance, including one (968-A) collected from a woods near the Arboretum in 1884, but we also hope to add more wild-collected accessions of this beautiful native tree as part of our ongoing Campaign for the Living Collections. Nancy Rose is the editor of Arnoldia. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23454","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14ea728.jpg","title":"2017-74-4","volume":74,"issue_number":4,"year":2017,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Rise and Fall of the Ornamental Callery Pear Tree","article_sequence":1,"start_page":2,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25617","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14eab6b.jpg","volume":74,"issue_number":3,"year":2017,"series":null,"season":null,"authors":"Culley, Theresa M.","article_content":"ARCHIVES OF THE ARNOLD ARBORETUM The Rise and Fall of the Ornamental Callery Pear Tree Theresa M. Culley O ne of the most notable heralds of spring in the eastern United States is the profuse blooming of ornamental pear trees in front yards and along city streets. The Callery pear (Pyrus calleryana), and particularly its many cultivars such as `Bradford', `Cleveland Select', and `Aristocrat', has become one of the most popular ornamental trees in North America. However, its commercial success has now become overshadowed by its tendency to spread along roadways and into natural areas through reseeding. Today this tree is considered invasive in many states, in stark contrast to how it grows in its native range in Asia. How did this tree become the scourge of land managers across North America? What has led to its fall from grace? To understand this fascinating story, we need to start at the beginning. Seeds From China Plant collector Frank N. Meyer in China in 1908. Toward the end of the nineteenth century, farming began to replace ranching in the western United States and there was a growing demand for improved crops that could thrive there. The United States Department of Agriculture (USDA) began to focus on importing new plants for testing and, in 1898, created the Foreign Seed and Plant Introduction Office. The mission of this office, headed by David Fairchild, was to locate and import economically important plants from other regions of the world. Fairchild was especially interested in China, which was thought to possess a wealth of unexplored botanical resources. Chinese plants were also expected to grow well in the United States because China's climate is very similar to that in the United States. In the early 1900s, Fairchild began searching for plant explorers who had the dedication and stamina to tolerate the physical discomforts and social isolation of travelling for months in distant lands. He found Frank N. Meyer (1875? 1918), a Dutch immigrant and former gardener who had a deep fascination with plants and saw nothing unusual about walking hundreds of miles on a botanical foray. Meyer eventually spent over ten years traveling across Asia looking for useful and valuable plants, seeking, in his own words, to \"skim the earth in search of things good for man.\" He eventually sent hundreds of shipments of cuttings and thousands of pounds of seeds back to the USDA. Many agricultural crops grown in the United States today, including certain grains, legumes, and fruits, resulted from Meyer's collections. But before he began his first Chinese expedition in 1905, Meyer visited several United States gardens to become familiar with Chi- ARCHIVES OF THE ARNOLD ARBORETUM Pyrus calleryana 3 The search for new plants from other countries became more urgent in the early 1900s when valuable orchards of the edible French pear (Pyrus communis) were being decimated by fire blight in the Pacific Northwest. This bacterial disease blackened leaves and branch tips of infected trees as if they were scorched by fire, eventually killing large fruit trees, and it was quickly spreading throughout the region. In the hopes of breeding resistance to fire blight into P. communis, Professor Frank C. Reimer of the Southern Oregon Experiment Station hastily began testing all available Pyrus species and varieties for resistance to this devastating disease. The initial results proved disappointing, so a call was put out to locate Pyrus species in Ernest H. Wilson made this photo of a 40-foot-tall Pyrus calleryana tree in a Chinese botanical garden on April 7, 1909. nese plants in their collections. He was most impressed with the Arnold Arboretum's collection and he met director Charles Sprague Sargent, who was keenly interested in Meyer's travels but who also had a complex and often difficult relationship with Fairchild. Meyer clearly respected the Arnold Arboretum--he once requested that his Chinese material be sent there instead of to a USDA station because he felt it would receive better care. However, Meyer's relationship with the Arnold's plant explorer Ernest H. Wilson was somewhat uneasy at first, likely because they first viewed each other as competitors since Sargent had also asked Meyer to collect for the Arboretum during his Chinese expeditions. While Meyer was willing to oblige, this arrangement would sometimes place Meyer in an awkward situation because Sargent's emphasis on capturing the diversity of the Chinese flora was often at odds with Fairchild's directive to focus only on economically important species. This 1890 USDA illustration shows a stem of the edible pear (Pyrus communis) cultivar `Le Conte' that had been inoculated with fire blight bacteria twelve days prior. The terminal \"shepherd's crook\" and blackened leaves are characteristic of fire blight. From the USDA Pomological Watercolor Collection, National Agricultural Library, Special Collections. Pyrus calleryana is simply a marvel. One finds it growing under all sorts of conditions; one time on dry, sterile mountain slopes; then again with its roots in standing water at the edge of a pond; sometimes in open pine forest, then again among scrub on blue-stone ledges in the burning sun; sometimes in low bamboo-jungle ... and then again along the course of a fast flowing mountain stream or in the occasionally burned-over slope of a pebbly hill. The tree is nowhere found in groves; always as scattered specimens, and but very few large trees were seen. In 1917, Reimer himself joined Meyer in China and they traveled together for several days, with Meyer showing Reimer the locations of Pyrus calleryana trees he had found. In his report, Reimer's amazement at this plant is evident, which also heralded his eventual emphasis on the species as rootstock: In its ability to endure diverse and adverse soil conditions, this species certainly is a marvel ... I found it growing in all the various soil types ... Flowers, leaves, and fruits of Pyrus calleryana collected in China and photographed by Frank Meyer in April 1917. Meyer's description of the photo from his South China Exploration typescript: \"Pyrus calleryana, natural size. A somewhat small-flowering type of a wild Calleryana pear, with rather tomentose foliage, which isn't full grown yet. Three fruits of last year's crop has persisted on the tree during the whole winter and spring. Note the very small size, on which account the Chinese call it the \"T'ang li\" or crab-apple pear, as these small fruits, with deciduous calyx, resemble the tiny apples of Malus spectabilis and M. baccata to a surprising degree.\" USDA NATIONAL AGRICULTURAL LIBRARY, SPECIAL COLLECTIONS other parts of the world that might be fire blight resistant. Many plant explorers, including E. H. Wilson and Emil Bretschneider, traveled to Asia in the early 1900s, in part to locate new Pyrus species, often with the material sent back to the Arnold Arboretum. In 1908, Wilson first introduced P. calleryana into the United States with several seed lots accessioned and grown at the Arnold Arboretum. In 1916, between his Chinese expeditions, Meyer visited the Pacific Northwest where he saw for himself the extensive fire blight destruction. He now understood the importance of his work because he learned from Reimer that resistance had only been found in the wild Chinese pear species P. calleryana and P. ussuriensis. However, Reimer needed more material for testing and Meyer agreed to collect and send back over 100 pounds of wild P. calleryana seeds during his next expedition. This was no small task since 25 pounds of cleaned seeds required at least 5,000 pounds of fruit. During his subsequent months in China, Meyer focused much of his effort on Pyrus calleryana. He painstakingly collected thousands of the marble-sized pear fruits in the field or bought them directly from local Chinese. He later wrote Fairchild that: USDA NATIONAL AGRICULTURAL LIBRARY, SPECIAL COLLECTIONS 4 Arnoldia 74\/3 ? February 2017 Frank Meyer photographed the environmentally adaptable Pyrus calleryana growing in a number of distinct habitats in China including along waterways with roots in standing water, in crevices in shale rock, within a dense jungle of bamboo, and, seen here, in shrubby, dwarfed form on a dry mountain top. ranging from heavy clays to light sandy soils and disintegrated rock. I found it growing in shallow ponds, along streams, well-drained moist loams, and on very dry poor hillsides and hilltops. In places it was observed where the layers of soil above the bedrock was not more than eight inches deep. THE ARNOLD ARBORETUM Pyrus calleryana 5 Upon his return, Reimer continued to work with other plant explorers to obtain Pyrus calleryana seeds for further testing. Tragically, Meyer never returned to the United States, drowning in the Yangtze River in late 1918 just as he was beginning his trip home. However, his much-anticipated collection of P. calleryana seeds was shipped back in his absence, to complete the task that he had begun years earlier. It is from many of these seeds that our story continues. `Bradford', the First Callery Pear Cultivar Over the next few years, Chinese seeds collected by Meyer and Reimer were planted in large numbers--primarily at the USDA Plant Introduction Stations at Corvallis, Oregon, and Glenn Dale, Maryland--to test their resistance to fire blight. Over time, interest in the species turned to its ability to serve as rootstock for the economically valuable edible French pear. For example, Reimer also saw that under favorable conditions in China, the tree \"is a rapid, vigorous grower, has a long growing season, and its leaves remain green and lusty until very late in the fall.\" In central China, the trees were often cut off for firewood every few years but they would put out \"new sprouts from the stumps and continue to live for many years.\" These wild trees also typically produced prominent thorns (actually sharp spur shoots) that effectively protected against herbivory. The species seemed to be adapted to mild climates, with Reimer suggesting, \"It is quite probable that it will not endure very severe winter climates.\" However, he also wrote that trees that originated from China had proved to be very hardy at the Arnold Arboretum over 10 years, despite the more severe winters near Boston compared to the native range in China. THERESA M. CULLEY Herbarium specimen of P. calleryana growing at the Arnold Arboretum in May 1918. The tree was grown from seeds collected in China by Ernest H. Wilson in April 1908. \"Thorns\" are naturally produced by both Callery pear trees in China and wild trees in the United States. This structure is technically a spur, a short, pointed shoot bearing leaves or flowers. Most Callery pear cultivars were selected in part for thornlessness. THERESA M. CULLEY THERESA M. CULLEY 6 Arnoldia 74\/3 ? February 2017 for resistance to fire blight, overall vigor, and stock-scion compatibility with P. communis. In 1952, one of the remaining 33-year-old trees of Pyrus calleryana from Meyer's Chinese seeds that was still growing near the Plant Introduction Station in Glenn Dale caught the eye of John Creech of the USDA (Creech 1973). This tree had thick, glossy leaves and an attractive globular form, with a lack of sharp spurs so typical of the species. Recognizing its potential as a landscaping tree, Creech grafted scions of it onto P. calleryana rootstock. This method of propagation means that every tree is genetically identical to the original mother tree. Creech named this cultivar `BradA typical `Bradford' Callery pear flowering in early spring. This tree has already lost a limb and the homeowner has tied straps around the inside ford', in honor of F. C. Bradford, branches in an attempt to prevent further breakage. the former horticulturist in charge of the Glenn Dale USDA station (Whitehouse et al. 1963). (Incidentally, the original `Bradford' tree was destroyed years later to make way for a parking lot). In 1954, Creech planted two-yearold `Bradford' clones in a nearby residential subdivision in University Park, Maryland, for a street tree study. `Bradford' swiftly became quite popular for its rapid growth, attractive foliage that was retained into late fall, extremely showy and abundant flowers in early spring, and its overall hardiness. The cultivar was commercially released around 1961 and then planted widely across the eastern United States in residential areas. However, by the early 1980s probA `Bradford' tree in West Chester, Ohio, that split during the winds of Hurlems with `Bradford' pears began ricane Ike in September 2008. to appear, especially a tendency for Reimer wrote that \"thousands of seedlings older trees to break apart during windstorms have been grown\" in the Pacific Northwest or under heavy snow loads. Its branching structure was to blame, as described by horticulturfrom Meyer's original seed \"to test this species ist Michael Dirr (1998), \" `Bradford' tends to thoroughly as a stock for our cultivated varieties.\" In Glenn Dale, Meyer's seeds were also develop rather tight crotches and I have seen planted out in large numbers to test the plants trees that were literally split in half ... the plant More Callery Pear Cultivars Appear Over the next few decades, additional P. calleryana cultivars were quickly introduced as improved replacements for older `Bradford' trees that had begun to split. For example, `Whitehouse', a narrow columnar form with a strong central leader, was selected in 1969 from seedlings still growing near the Plant Introduction Station in Glenn Dale. This cultivar presumably began as a seed from the original `Bradford' tree that had been pollinated by one of Fall foliage color on Callery pear cultivars and seedlings ranges from yellow the other wild P. calleryana plants at to bright red and purple. the station. Similarly, `Redspire' also arose as a `Bradford' seedling and was patented in 1975. At the opposite side of the country, `Autumn Blaze' was selected in 1969 from among several hundred seedlings of P. calleryana growing at the rootstock research nursery in Corvallis. This cultivar, known for its striking red to purple leaf coloration in the fall, originated from seeds that Reimer had brought back from China during his trip with Meyer in 1917 or later in 1919. Other cultivars such as `Avery Park' and `Grant St. Yellow' also originated in the late 1960s and 1970s near Corvallis, most likely from Chinese seeds imported into that area. Callery pear cultivars became extremely popular as urban street trees in the Over time, additional cultivars eastern United States. arose from other areas of the country, presumably from different seed sources. These and other Callery pear cultivars became For example, `Aristocrat' arose in Indepenexceedingly popular as ornamental landscapdence, Kentucky, while the cultivars `Cleveland ing trees for residential and commercial use. Select', `Chanticleer', and `Stone Hill' were all Not only were they beautiful, fast growing, and derived from the same street tree in Cleveland, inexpensive, but they were also extremely tolerant of very difficult growing conditions. In Ohio. The cultivar `Valzam' was found growing commercial areas, for example, the tree could among `Cleveland Select' trees in Perry, Ohio in thrive in the harsh conditions of parking lot 1975--most likely an offspring of that cultivar islands and between streets and sidewalks, but with unknown paternity. THERESA M. CULLEY will ... fall apart because of the development of many branches around a common length of the trunk.\" LESLIE J. MEHRHOFF, UNIVERSITY OF CONNECTICUT, BUGWOOD.ORG Pyrus calleryana 7 8 Arnoldia 74\/3 ? February 2017 where temperatures were excessively high and water was scarce. Following the tragic events of September 11, 2001, when the World Trade Center's twin towers fell in New York City, a Callery pear tree at the site was found still alive but severely burned with damaged roots and branches. Known today as the \"Survivor Tree,\" it was rescued from the site, taken to a local nursery to recover, and later replanted back in the memorial park at Ground Zero as a symbol of resilience. In 2005, `Chanticleer' was chosen as the Urban Tree of the Year by the Society of Municipal Arborists, who noted that, \"This tree has all of the character and quality of a sheared topiary specimen plus, of course, the magnificence of its spring, summer, and fall outer garments-- the white flower, crisp glossy green summer foliage, and full fall color.\" Some homeowners associations in areas of the United States even had a requirement that a specific Callery pear cultivar had to be planted in each front yard. In fact, the Callery pear had become so popular that Michael Dirr lamented in 1989 that \"cookie-cutter Bradfords ... inhabit almost every city and town to some degree or another; the tree has reached epidemic proportions and is over-planted.\" In 2009 alone, the species generated over $23 million in sales across the country (USDA 2010), including continuing sales of `Bradford'. Although the majority of commercial sales occurred in the eastern and southern United States, cultivars were also available along the western coastal states. The tree had reached its epitome of fame and glory. The Fall From Grace As `Bradford' and other Callery pear cultivars surged in popularity, early indications of problems began to appear. Pyrus calleryana had escaped cultivation as early as 1964 in Arkansas and 1965 in Maryland (Vincent 2005), but it was not until the 1990s that the species began to be more widely noticed in natural areas, especially in southern states. For example, Michael Vincent of Miami University (Ohio) examined 300 P. calleryana herbarium specimens collected across the nation beginning in 1964. He found that 1% of all specimens were collected in each of the periods 1964?1969 and 1970?1979 before a dramatic increase began in 1980?1989 (17% of all specimens), continuing through 1990?1999 (31%), and mounting rapidly in the last three years of the study, 2000?2003 (50%). By the late 1990s, members of several Internet gardening forums began noting the increasing numbers of wild pears beside roadways along the midAtlantic coast, largely in the Maryland area. By the middle of the twenty-first century's first decade, thousands of young wild pear seedlings were growing undetected in the roadside vegetation across the southern and eastern United States. But as they began to flower in their third or later year, their profuse early spring blooms started to give them away. As each successive year revealed more and more wild pears blooming, public alarm began to sound. Land managers began to notice wild pears appearing in all types of habitats--along forest edges, in wetland areas, and even within forests. This is not surprising given Meyer's and Reimer's remarks about the many different habitats where P. calleryana is found in its native range. In the United States, word eventually began to spread of not only the slippery mess caused by pear fruits littering sidewalks, the difficulty in removing dense stands of thorny trees in natural areas, and the putrid smell of the flowers, but also increasing concern of liability caused by falling tree limbs damaging property and injuring people. Pyrus calleryana (often indicated as just \"Bradford pear\") began to appear on watch lists and invasive plant lists in several eastern and southern states. But why had this pear, which had behaved for decades as a popular landscaping tree, suddenly start to spread uncontrollably? The answer lies in the reproductive system of the species as well as its horticultural history. As with most other pears, Pyrus calleryana has a genetically controlled self-incompatibility system that prevents individual trees from pollinating themselves, thus requiring outcrossing among unrelated individuals. When `Bradford' was first introduced and became so wildly popular, `Bradford' trees were unable to cross-pollinate (since they were all genetically identical) and fruits were never produced. As additional cultivars were introduced they were often commercially marketed as \"self-sterile\" JOE BOGGS Pyrus calleryana 9 THERESA M. CULLEY Invasive population of wild Callery pears grows in a field next to Interstate I-75 in Butler County, Ohio. When two or more Callery pear cultivars are planted nearby, abundant fruits are usually produced. or even \"seedless\"--this was true, as long as each cultivar was grown in isolation. Cultivar patents and promotional material included notes such as \"[fruit set] very low (about 5 to 10%), usually only one fruit per cluster\" (`Autumn Blaze'); \"little or no fruit and the fruit that is produced is small and hard\" (`Trinity'); and \"self-sterile\" with fruits typically not abundant and only produced when \"planted near another clone\"(`Aristocrat'). The last point is the lynchpin of this story. Although each Callery pear cultivar cannot produce fruits on its own, fruits can easily develop when two or more cultivars--which are genetically different and therefore cross compatible--are planted together (Culley and Hardiman 2007). Cross-pollination is promoted by insect pollinators, especially bees, which frequently fly over a mile each day visiting all flowers within their range. This has a large impact on the magnitude of the pear problem. For example, if a large residential area only contains `Bradford' trees, no fruits would be formed. But if a new resident moves in and plants a single `Aristocrat' in her yard, that new tree now has the potential to cross-pollinate all the `Bradford' trees within a mile-wide range, and vice-versa. This could trigger a sudden outburst of fruit within a single year. Such massive fruiting may even go undetected at first because some people expect any pear, including P. calleryana, to produce large edible fruits like a `Bartlett' pear, and they do not recognize the small fruits of Callery pear. During the winter months, these fruits are consumed by birds that then defecate the seeds as they fly or roost in trees or along power lines, thereby spreading the species into new areas. In fact, genetic analysis of new wild Callery pear populations have confirmed that wild plants are typically F1 hybrids of cultivars planted in the surrounding residential and commercial areas (Culley and Hardiman 2009). In older populations, such as near the Glenn Dale station where `Bradford' was first discovered, wild pear populations largely consist of advanced generation hybrids. Cross pollination between mature specimens of Callery pear cultivars is not the only way fruit production can occur. To maintain their genetic identity, Callery pear cultivars are clonally produced by vegetative propagation. Stem cuttings of Callery pears are difficult to root so most trees are propagated for commercial sale by grafting. In this process, the scion material of the selected cultivar is grafted onto THERESA M. CULLEY LESLIE J. MEHRHOFF, UNIVERSITY OF CONNECTICUT, BUGWOOD.ORG 10 Arnoldia 74\/3 ? February 2017 to come from a landscape planting of multiple cultivars or even from a single grafted tree. In a genetic parentage study of a wild Callery pear population in southwestern Ohio, it was discovered that at least 17% of wild trees had a rootstock parent (Culley et al. 2011). The Future The introduced range of Pyrus calleryana in the United States is currently restricted by the species' limited cold tolerance, as predicted decades ago by Reimer and Meyer. However, hardier cultivars are now being developed that will expand A flock of European starlings (Sturnus vulgaris) eats Callery pear fruits in Callery pear's landscape presence. a parking lot. In addition, wild Callery pear is expected to continue to spread northward as global climate change causes shifts in warmer temperatures. In fact, wild Callery pears have already been observed around Madison, Wisconsin (USDA Hardiness Zone 5a), an area where they were thought never to survive. What can be done to prevent the continued spread of the wild Callery pear? First, Callery pear cultivars need to be carefully phased out of commercial production and replaced with suitable alternatives. The latter is critical, as it would allow plant breeders and the nursery industry to recoup any economic loss and remain profitable. In fact, sterile varieties of P. calleryana are curThe rootstock of commercially grafted Callery pear trees can occasionally rently being developed. Second, as sprout. If left to grow and flower, the rootstock can cross-pollinate the upper cultivated trees within the landscape scion of the tree. break apart or decline they should be replaced with these alternatives. This is rootstock, which is usually P. calleryana seedsomething already happening in many towns lings. The two sections grow together, resulting nationwide, as Callery pears are being replaced in a tree composed of two genotypes. Occasionwith different tree species. Finally, if homeally, the rootstock of a planted Callery pear owners choose to keep cultivated Callery pears cultivar may develop shoots that eventually growing in their yards, they must take responsiflower. In such a case, the rootstock now has bility for ensuring that fruits are not produced. the potential to cross-pollinate the upper scion Callery pear fruit production can be reduced of the same tree, triggering fruit production. So as much as 95% by spraying flowering trees it is possible for a wild population of pear trees Pyrus calleryana 11 with ethephon, a plant growth regulator that does not affect the flowers' appearance but does make them incapable of developing into fruit. But even if all these suggestions could be accomplished, the sad reality is that wild Callery pear will continue to be a growing problem in the years to come because so many cultivars and their rootstock are already established in the landscape. Today, the Callery pear story is another example of how even the best of human intentions can go awry. The pear was introduced into the United States for the best of reasons-- to save the valuable crop of P. communis on the West Coast from fire blight in the 1920s. Decades later, `Bradford' and other Callery pear cultivars were selected and promoted to give gardeners and landscaping professionals additional highly tolerant and attractive trees for the landscape. These were all good and sensible ideas at the time, especially since the majority of introduced species in the United States never become invasive. The resulting spread of wild P. calleryana into the American landscape was unanticipated and completely unintentional. The best that we can do today is to view the Callery pear as a lesson on the importance of considering how mixes of ornamental cultivars may contribute to invasive spread of certain species. By learning from our past history, we can better understand why certain species become invasive, and thus we can work more effectively to prevent invasive spread of species in the future. Acknowledgments The author would like to thank the many researchers, land managers, nursery professionals, and students who were invaluable in uncovering this story. Most notably, this project was sparked by Marjie Becus, who first pointed out wild Callery pear to the author during their vegetational rambles, and Robert Naczi, who mentioned the Internet discussion on the topic shortly after. The author also thanks Joe Boggs and Michael Vincent for their willingness to share their thoughts and expertise, as well as Larissa Glasser for locating Frank Reimer's original notes in the Arnold Library. This article is dedicated to the memory of Sarah Elizabeth Reichard, who was instrumental in highlighting the importance of horticultural introductions in plant invasions as well as the necessity of working with nursery interests to create practical and effective solutions. She will be missed. References Creech, J. L. 1973. Ornamental Plant Introduction-- Building On the Past. Arnoldia 33(1): 13?25. Culley, T. M. and N. A. Hardiman. 2007. The beginning of a new invasive plant: A history of the ornamental Callery Pear tree in the United States. BioScience 57: 956?964. Culley, T. M. and N. A. Hardiman. 2009. The role of intraspecific hybridization in the evolution of invasiveness: A case study of the ornamental pear tree Pyrus calleryana. Biological Invasions 11: 1107?1119. Culley, T. M., N. A. Hardiman, and J. Hawks. 2011. The role of horticulture in plant invasions: how grafting in cultivars of Callery pear (Pyrus calleryana) can facilitate spread into natural areas. Biological Invasions 13: 739?746. Cunningham, I. S. 1984. Frank N. Meyer: Plant Hunter in Asia. Ames, Iowa: The Iowa State University Press. Cunningham, I. S. 1984. Frank Meyer, Agricultural Explorer. Arnoldia 44(3): 3?26. Dirr, M. A. 1998. Manual of Woody Landscape Plants: Their Identification, Ornamental Characteristics, Culture, Propagation and Uses. Fifth Edition. Champaign, Illinois: Stipes Publishing L.L.C. Meyer, F. N. 1918. South China Explorations: Typescript, July 25, 1916?Setpember 21, 1918. The National Agricultural Library. Available online at: https:\/\/archive.org\/details\/ CAT10662165MeyerSouthChinaExplorations Reimer, F. C. (undated) Report of a Trip to the Orient to Collect and Study Oriental Pears. The Arnold Arboretum. United States Department of Agriculture. 2010. Census of Horticultural Species (2009). Volume 3, Part 3. Available online at: https:\/\/www.agcensus.usda. gov\/Publications\/2007\/Online_Highlights\/ Census_of_Horticulture_Specialties\/index.php Vincent, M. A. 2005. On the Spread and Current Distribution of Pyrus calleryana in the United States. Castanea 70: 20?31. Whitehouse, W.E., J.L. Creech, and G.A. Seaton (1963) Bradford Ornamental Pear--A Promising Shade Tree. American Nurseryman 117: 7?8, 56?61. Theresa Culley is a Professor of Biological Sciences at the University of Cincinnati. She also serves as a board member of the Midwest Invasive Plant Network and is a past president of the Ohio Invasive Plants Council (OIPC). She currently is the Chair of the OIPC's Invasive Plant Assessment Team for the state of Ohio. "},{"has_event_date":0,"type":"arnoldia","title":"Plant Dye Identification in Japanese Woodblock Prints","article_sequence":2,"start_page":12,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25616","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14eab26.jpg","volume":74,"issue_number":3,"year":2017,"series":null,"season":null,"authors":"Newman, Richard; Wright, Joan; Derrick, Michele","article_content":"Plant Dye Identification in Japanese Woodblock Prints Michele Derrick, Joan Wright, Richard Newman W oodblock prints were first produced in Japan during the sixth to eighth century but it was not until the Edo period (1603?1868) that the full potential of woodblock printing as a means to create popular imagery for mass consumption developed. Known broadly as ukiyo-e, meaning \"pictures of the floating world,\" these prints depicted Kabuki actors, beautiful women, scenes from history or legend, views of Edo, landscapes, and erotica. Prints and printed books, with or without illustrations, became an integral part of daily life during this time of peace and stability. Prints produced from about the 1650s through the 1740s were printed in black line, sometimes with hand-applied color (see figure 1). These colors were predominantly mineral (inorganic) pigments supplemented by plant-based (organic) colorants. Since adding colors to a print by hand was costly and slowed production, the block carvers eventually hit upon a means to create a multicolor print using blocks that contained an \"L\" shaped groove carved into the corner and a straight groove carved further up its side in order to align the paper to be printed (see figure Figure 1. Actors Sanj Kantar II and Ichimura Takenoj IV, (MFA 11.13273), 2). These guides, called kento, are located about 1719 (Kyho 4), designed by Torii Kiyotada I, and published by in the same location on each block. They Komatsuya (31.1 x 15.3 cm). Example of a beni-e Japanese woodblock ensure consistent alignment as each color print with hand-applied color commonly made from the 1650s to 1740s. is printed onto a single sheet of paper. From the 1740s to about 1765, the first block green. From 1765 on, the skills required to use printed colors appeared on simple two- or threethe kento registration system reached a level color images (see figure 3). These benizuri-e where several color blocks could be expertly (\"red pictures\") utilized red, blue, or yellow; printed and full-color nishiki-e or \"brocade sometimes these colors were over-printed to prints\" such as those designed by Suzuki Harunobu (1725?1770) became the standard create the secondary colors purple, orange, and Plant Dye Identification 13 Figure 2. A Japanese woodblock that illustrates the `L' shaped kento groove added in the corners of each block to aid in the alignment of the paper for printing multiple colors. (see figure 4). Vibrant full-color prints designed by well-known artists such as Torii Kiyonaga (1752?1815) and Kitagawa Utamaro (1753?1806), produced and marketed by the great publishing houses of Tsutaya Juzaburo and Nishimura Yohachi, defined the period from 1781?1801, which is often referred to as the Golden Age of the Japanese woodblock print (see figures 5 and 6). Figure 3. Actor Ichikawa Danz III as Adachi Hachir, (MFA 11.19030), 1762 (H?reki 12) 11th month, designed by Torii Kiyomitsu I, and published by Urokogataya Magobei (30.2 x 14.2 cm). Example of a benizuri-e Japanese woodblock print with a 3-color palette commonly made from the 1740s to 1765. 14 Arnoldia 74\/3 ? February 2017 tures, and seals on the print into the Museum System collections database (TMS), enabling our current research as well as numerous exhibitions and publications. Of additional significance, the collection holds a number of multiple impressions of a single image, thus providing an ideal setting to identify, survey, and understand the organic and inorganic colorants used in traditional Japanese color woodblock printing. In 2013, the Asian Conservation and Scientific Research divisions began a large-scale survey of the colorants used in the MFA collection of Japanese prints. This effort uses only nondestructive techniques, which means no samples are required of the prints that are formed by minimal levels of colorants absorbed into their paper fibers. The first two techniques used for this study are standard methods used in museum labs: X-ray Fluorescence (XRF) and Fiber Optic Reflectance Spectroscopy (FORS). A new and previously little-used technique of Excitation-Emission Matrix (EEM), or 3D, fluorescence spectroscopy was also used to successfully characterize several additional dyes. Colorants The plant-based red, yellow, and blue dyes long considered to make up the palette of Japanese woodblock prints are summarized in Table 1. The list is based on various published sources, including early Japanese literature and analytical studies, and may not be comprehensive. The inorganic pigments used in the prints, such as red lead, hematite, and orpiment, can be easily estimated by XRF (see Table 2). The traditional organic blues, dayflower and indigo, can be confidently identified by FORS in the visible and near-infrared ranges. Both XRF and FORS were used in the examination of the prints discussed in this article. The unique component of this study was the use of EEM fluorescence spectroscopy to identify the yellow and red natural Figure 4. Courtesan Watching Two Kamuro Make a Snow Dog, (MFA 21.4463), about 1767?68 (Meiwa 4?5), designed by Suzuki Harunobu (28.5 x 21.8 cm). Example of full color printing, nishiki-e, characteristic of the early years from 1766?1780. Japanese Woodblock Prints at the MFA The Museum of Fine Arts (MFA), Boston, holds a collection of over 50,000 Japanese woodblock prints and illustrated books. This represents the largest number of such art works in a single location outside of Japan. In 2010, a major five-year project to accession, image, and re-house this vast collection of Japanese woodblock prints was completed with cataloguing ongoing. This project placed information about each print along with its image and the translation of any Japanese text, signa- Plant Dye Identification 15 Mass Spectrometer detector (LC\/MS). Afterward, each material was prepared by historical methods, then printed onto Japanese paper. For the materials listed in Table 1, all were available from documented sources as raw materials, except for Toringo crabapple (Malus sieboldii) and Coptis japonica, Japanese goldthread, a member of the buttercup family. Fortunately, the Arnold Arboretum generously supplied samples of branches from five Malus sieboldii specimens in their collection. In lieu of C. japonica, a sample of threeleaf goldthread (Coptis trifolia), a related species native to North America, was obtained from a biologist in Vermont. Results An initial set of 213 Japanese woodblock prints were examined at the MFA by the combined techniques of XRF, FORS, and EEM fluorescence. These prints covered the time period 1700 to 1800 (see Table 5). The goal for the analysis of the prints was to obtain an overview of the colorants used by artists active in each time period; this goal was later expanded to include information on publishers, since they were probably responsible for the final colorant decisions. Our research is ongoing and is conducted as time allows. It is hoped that the analysis Figure 5. Actors Matsumoto Kshir IV as Ukita Sakingo and Sawamura Sjr III of a more extensive set of prints will as the Ghost of Takao, with chanters Tomimoto Itsukiday? and Tomimoto Awatay?, provide definitive information on the and accompanist Sasaki Ichishir?, (MFA 11.13921), 1788 (Tenmei 8) autumn, relationships between colorants, pubdesigned by Torii Kiyonaga and published by Nishimuraya Yohachi (38.8 x 26.8 lisher, artist, and period. (See Table 6 cm). Example of full color printing, nishiki-e, characteristic of Japanese woodfor the results from the example prints block prints made from 1781 to 1801. mentioned in this article.) organic colorants on the prints. Examples of Even with this limited data set, several patterns of colorant use were consistently found the color and line contour maps obtained as for the eighteenth century time period. From results for the fluorescence analysis are shown the beginning, it was clear that the prints often in Tables 3 and 4. contained more than one yellow, red, or blue For this project, it was important to obtain colorant. Though it seems logical, since each reference materials from documented sources colorant has unique tonal properties, this findfor each of the materials to determine the best ing was significant in terms of analysis, indimethod for its identification. Once the reference samples were obtained, they were authencating it was imperative to analyze multiple ticated using Liquid Chromatography with a colored regions for each print. Because of the 16 Arnoldia 74\/3 ? February 2017 PLANT SOURCE JAPANESE NAME COMMON ENGLISH NAMES PART OF PLANT USED COLOR Caesalpinia sappan suo, suwo sappanwood heartwood red Carthamus tinctorius benibana safflower florets separated from capitulum red Rubia akane akane Japanese madder roots, stems red Rubia tinctorum seiyo-akane European madder roots, stems red Berberis thunbergii megi Japanese barberry roots, stems yellow Coptis japonica; C. trifolia woren goldthread roots, stems yellow Curcuma longa (syn. C. domestica); C. aromatica ukon turmeric rhizomes yellow Gardenia jasminoides (syn. G. augusta) kuchinashi gardenia juice or extract from fruit yellow Garcinia hanburyi, G. morella shio, te-o; kusa shio gamboge resin\/powder yellow Malus sieboldii (syn. Pyrus toringo) zumi Toringo crabapple bark yellow Miscanthus tinctorius; M. sinensis kariyasu silver grass grass cut when flowering spikes form, then dried over the winter yellow Myrica rubra yama-momo mountain peach, red bayberry bark yellow Nandina domestica nanten nandina, heavenly bamboo branch yellow Phellodendron amurense kihada Amur corktree inner bark of trunk yellow Styphnolobium japonicum (syn. Sophora japonica) enju Japanese pagoda tree unopened flower buds yellow Table 1. Common Asian natural red and yellow organic colorants in Japanese woodblock prints. expansion to the use of mixtures and colorant variations from 1781 to 1801, it was common to find three types of yellow, two reds, and two blues in a single print (example figure 5). Thus, while 213 prints were studied, there were over 1,500 individual analysis points. As expected from the literature, safflower (Carthamus tinctorius) was the primary red and pink colorant used consistently for all of the time periods and methods of application. Surprisingly, however, the second most prolifically used red was madder. While the analytical methods used in these tests, could not distinguish between Japanese madder (Rubia akane) and European madder (Rubia tinctorum), one or both of these colorants were consistently found Plant Dye Identification 17 on prints in all four of the described periods with their use increasing from 20% up to 50% over the hundred-year period. The yellow colorants changed significantly over the hundred-year period from the sole use of flavonoids and gamboge during the beni-e hand-applied color period (1710?1740s) to the predominant use of turmeric and orpiment (an arsenic sulfide mineral) for the elaborate designs and techniques used for full color printing from 1781?1801. Inorganic pigments were found on most prints examined for each time period. While the use of orpiment (As2S3) increased significantly, the use of hematite (an iron oxide, Fe2O3) and lead (Pb) were constant. Other inorganic pigments were occasionally found, such as added decorations using ground metallic brass (figure 1). Vermilion, a mercurycontaining pigment commonly used for paintings, has been mentioned as a definitive colorant in Japanese printing. However, this study found only two prints containing vermilion, indicating it may not have been commonly used. Of interest to us were the compositions for purples and greens. Mixtures or overprinting transparent colors were noted in many prints from the 1740s on. In all analyzed purple regions, our results showed mixtures of safflower and dayflower (see figures 4 and 5). The presence of this mix- Figure 6. The Heron Maiden (Sagi musume) from the series An Array of Dancing Girls of the Present Day, (MFA 11.14364), 1793?94, designed by Kitagawa Utamaro I and published by Tsutaya Jzabur (Kshod). MINERAL SOURCE JAPANESE NAME COMMON ENGLISH NAMES CHEMICAL FORMULA COLOR hematite benigara red ocher Fe2O3 red red lead tan, entan red lead Pb3O4 red vermilion shu, shin-sha vermilion HgS red goethite odo yellow ocher FeO(OH) yellow orpiment kio, sekio, shio orpiment As2S3 yellow azurite iwagunjo azurite Cu3 (CO3 )2 (OH)2 blue synthetic gunjo ultramarine Na4-8 Al 6 Si 6 O24 S2-4 blue Table 2: Common natural inorganic colorants used in Japanese woodblock prints. 18 Arnoldia 74\/3 ? February 2017 ture throughout the history of color printing seems to indicate that the tone obtained by mixing dayflower blue and safflower was preferred over other possible mixtures of reds and blues (for example, indigo and madder) to yield purple. The green regions varied more often, with earlier prints showing overprinting of turmeric with dayflower (figure 3) while later prints showed a more vibrant green made by mixing orpiment and indigo (figures 4 and 5). Two aspects of the results in this study seemed unusual. First, no examples of either gardenia (Gardenia jasminoides) or berberine colorants (e.g., Berberis thunbergii, Coptis japonica, Phellodendron amurense) were found in the analysis of 557 yellow spots in 213 prints. These plants, which generally grow in the highlands, have been described as common Chinese colorants that were used in the Japanese islands. The colorants were mentioned in the literature as being used for eighteenth century Japanese woodblock prints, but were not found on any prints analyzed in this preliminary study. Second, madder was found on 142 red analysis locations in 90 out of the 213 prints (42%). While madder was available in Japan and was European madder (seiko akane) On paper PMT=600 ex=550\/em=595 Japanese madder (akane) On paper PMT=675 ex=545\/em=585 E XC I TAT I O N WAV E L E N G T H (nm) Safflower (benibana) On paper PMT=630 ex=530\/em=570 Sappanwood (suo) On paper PMT=750 ex=560\/em=615 E M I S S I O N WAV E L E N G T H (nm) Table 3. Color and line contour plots for EEM fluorescent patterns of organic red Japanese colorants. Excitation and emission maxima are listed for the most intense spot. Plant Dye Identification 19 Turmeric (ukon) On paper PMT=600 ex=470\/em=525 Japanese pagoda tree (enju) On paper PMT=625 ex=450\/em=510 Gamboge (shio) On paper PMT=625 ex= -- \/em= -- E XC I TAT I O N WAV E L E N G T H (nm) Silver grass (kariyasu) On paper PMT=700 ex=455\/em=510 Amur corktree (kihada) On paper PMT=550 ex=445\/em=525 Gardenia (kuchinasi) On paper PMT=650 ex=350\/em=450 E M I S S I O N WAV E L E N G T H (nm) Table 4. Color and line contour plots for EEM fluorescent patterns of organic yellow Japanese colorants. Excitation and emission maxima are listed for the most intense spot. 20 Arnoldia 74\/3 ? February 2017 1650s?1740s 1740s?1765 Hand-colored prints, Beni-e and Urushi-e 16 Limited-color prints, Benizuri-e Orpiment (As) Red ocher (Fe) Red lead (Pb) Sappanwood Madder Safflower NUMBER OF PRINTS Gamboge WOODBLOCK PRINT STYLE Flavonoid DATE RANGE PERCENT OF PRINTS CONTAINING THAT COLOR WITHIN THAT STYLE Turmeric Table 5. Summary of red and yellow organic and inorganic colorants found on 213 Japanese woodblock prints. The results are given as the percentages (%) of the EEM fluorescent pattern type, or element (as determined by XRF), attributed to the prints of each style of production. More than one type of each color was often used within the same print. Over 1,500 points were analyzed in this group of prints. 0 75756319 6 1219 0 21 622419383314141014 1766?1780 Full-color prints, Nishiki-e: First Period 80 362823783926252050 1781?1801 Full-color prints, Nishiki-e: Golden Age 96 72 7 1 915111201471 used prolifically as a textile colorant, it has not been previously mentioned as a possible colorant for printing. Madder may have been used as a substitution for more costly reds such as safflower (benibana) in order to keep the market price of an individual print affordable. With most of the print collection in the MFA, it is not always known whether the prints are first editions or later runs. Thus, there is always the possibility that madder was used for later editions, even though the date of the print is listed based on its initial production. Further work will be done to compare impressions and examine the paper fibers and formation methods to clarify the time periods of the madder use. Additional Information on Specific Colorants Safflower: benibana The florets of safflower (Carthamus tinctorius) produce a wide range of colors from cherry red to pink (figure 7). Native to northern India and the Near East, this popular dye plant was widely cultivated throughout Asia and Europe by the end of the thirteenth century. It is a tender annual with spiny leaves and composite flower heads containing many yellow to orange disk florets. The florets are picked, then dried and crushed into a paste. The paste is washed with water to remove the non-lightfast yellow chromophores including several quinochalcones. The red colorant, primarily carthamin, is then extracted in an alkaline bath. The deepest reds are obtained through several initial washings to remove all of the water-soluble yellows. Red regions containing safflower were usually seen as bright fluorescence during the preliminary examination of the prints with a hand-held ultraviolet (UV) light. Thus, it was no surprise that the EEM fluorescence technique provided a unique and definitive pattern for safflower, even when it was visually observed as a faded brown tone. In addition to the fluorescence for the red chromophore, the pattern often contained an additional peak for the yellow chromophore that was supposedly removed in the preparation of the red colorant but often needed several washings for complete elimination. Printed examples of the safflower colorant can be seen in figures 1, 3, 4, and 5. In figure 1, the Kiyotada I hand-colored print from the beni-e period, safflower was found on the base of the umbrella and the flowers on the woman's kimono. Looking at the later Harunobu print from 1767?68 (figure 3), safflower was found on the purple robe of the kneeling child. In the 1788 print by Kiyonaga (figure 4), safflower was used for a pink collar, purple sleeve, and orange frame. Madder: akane and sieyo-akane The roots of madder plants (from Rubia tinctorum, Rubia akane, and many others) produce a deep true red color that was widely prized throughout the world (see figure 8). All madder family (Rubiaceae) plants give strong red dyes with the colorants concentrated in the parenchyma of the roots and stems, even though the plant and flowers do not exhibit any red colors. For processing, the roots were typically harvested in the autumn after a minimum of Plant Dye Identification 21 FIGURE MFA # ARTIST\/TITLE ANALYSIS POINTS ANALYSIS INTERPRETATION 1 11.13273 Torii Kiyotada I Blank - womans face red on umbrella . . . . . . . . . . . . pale robe man's chest . . . . . . . woman's robe near collar . . . . orange corner man's robe . . . . skirt yellow and black . . . . . . -- Safflower Flavonoid Flavonoid Gamboge Brass flakes* 3 11.1903 Torii Kiyomitsu I Blank - background -- green bell . . . . . . . . . . . . . . . . . Turmeric overprinted with dayflower** blue . . . . . . . . . . . . . . . . . . . . . Dayflower** yellow trim . . . . . . . . . . . . . . . Turmeric red foot . . . . . . . . . . . . . . . . . . Madder 4 21.4463 Suzuki Harunobu Blank- white snow -- yellow sky . . . . . . . . . . . . . . . . Flavonoid green bush . . . . . . . . . . . . . . . . Orpiment* mixed with indigo** red post . . . . . . . . . . . . . . . . . . Sappanwood red ribbon . . . . . . . . . . . . . . . . Sappanwood brown robe . . . . . . . . . . . . . . . Safflower mixed with dayflower** orange leaf on robe . . . . . . . . . Flavonoid mixed with sappanwood and safflower 5 11.13921 Torii Kiyonaga Blank - face -- red ground cloth . . . . . . . . . . . Madder yellow pants on musician . . . Flavonoid green grass . . . . . . . . . . . . . . . . Orpiment* mixed with indigo** green knee . . . . . . . . . . . . . . . . Orpiment* mixed with indigo** pink collar on woman . . . . . . Safflower pink shoulder on man . . . . . . Safflower orange scroll border . . . . . . . . Safflower mixed with turmeric purple cuff on man . . . . . . . . . Safflower mixed with dayflower** yellow at bottom . . . . . . . . . . Turmeric yellow fence . . . . . . . . . . . . . . Turmeric 6 11.14364 Kitagawa Utamaro I Blank - face center yellow flower . . . . . . . . green in hat . . . . . . . . . . . . . . . yellow ribbons . . . . . . . . . . . . pink kimono . . . . . . . . . . . . . . brown in hat . . . . . . . . . . . . . . blue on kimono . . . . . . . . . . . . -- Flavonoid Flavonoid with dayflower** Flavonoid Safflower Safflower Dayflower** Table 6. Summary of the analytical results for four Japanese woodblock prints covering the range of eighteenth century printing styles. Identification of colorants was made by EEM fluorescence, XRF*, and FORS**. KEITH LAWRENCE, MFA, BOSTON, HTTP:\/\/CAMEO.MFA.ORG\/WIKI\/SAFFLOWER 22 Arnoldia 74\/3 ? February 2017 Figure 7. Safflower (Carthamus tinctorius). Photomacrograph of dried florets for prepared dyes and an example of safflower dye on paper. two years growth. The plants were marketed as whole dried roots rather than as a powder. Madders, a general name applied to anthraquinone-containing dyes extracted from plants from various genera and species, often exhibit strong (orange) fluorescence in a work of art when examined under ultraviolet radiation. The strong fluorescence is usually stated to be associated with purpurin, a common anthraquinone found in many types of madder, including both Rubia tinctorum and R. akane, a plant native to Japan. It is not certain when R. tinctorum was first utilized in Japan, or from where it would have originated, but it does not appear to be possible to distinguish it from Rubia akane based on its EEM fluorescence pattern. Printed examples of the intense red madder colorant can be seen in figures 3, 5, and 6. Figure 3, the Kiyomitsu I limited color print from 1762, has madder as the sole red colorant and it was used for the man's robe, face, and feet. Looking at the later print by Kiyonaga (figure 5), madder was used for the red cloth under the musicians. When it was found, the madder EEM pattern was very distinct and its color was a bright deep red. Madder was not found in secondary colors such as purple or orange in the prints examined for this study. Red dyewood: suo The insoluble red dye from sappanwood (Caesalpinia sappan) and other types of red dyewoods (sandalwood, barwood, narrawood, padauk, camwood, Brazilwood, etc.) were prepared as colorants by pounding chips of the heartwood into a paste mixed with a little oil (see figure 9). These were formed into cakes or bars for storage and sale. The red colorant was so popular in the seventeenth and eighteenth centuries that many of these species are now extinct or endangered. Its color was said to be orange-red, brownish-red, or cinnamon-like. Using our references, the EEM spectra could easily distinguish the sappanwood fluorescence pattern from safflower and madder. However, the reference spectra for a few other types of red wood dyes, such as sandalwood and Brazilwood, produced similar but not identical fluorescence spectra. Thus, it was difficult, if not impossible, to differentiate between the various red dyewood sources. In this study, the red dyewood fluorescence pattern was not often found, its use being limited to just a few artists and publishers. The print by Harunobu (figure 4) shows an example of the red dyewood. It was used for both the red and the orange regions, while the Plant Dye Identification 23 purple colorant was found to contain safflower and dayflower. KEITH LAWRENCE, MFA, BOSTON, HTTP:\/\/CAMEO.MFA.ORG\/\/MADDER Turmeric: ukon Described as the most popular yellow colorant in the world, the rhizomes of turmeric (Curcuma longa) produce a bright yellow orange dye that is commonly used for food and textiles (see figure 10). Native to India, turmeric is now cultivated worldwide. Though a perennial herb, the plant is often completely harvested, then the roots are cooked, dried, and ground into a powder. Turmeric is a direct dye with high tinctorial strength that began its use as a fabric dye prior to the tenth century and is still used today as a curry seasoning. Yellow regions containing turmeric usually were brightly fluorescent during the preliminary examination of the prints with a hand-held UV light. The dye produced a very clear, consistent fluorescence pattern, likely because of its single primary chromophore. In this study, the printed examples that contain turmeric are figures 3 and 5. In the print by KEITH LAWRENCE, MFA, BOSTON, HTTP:\/\/CAMEO.MFA.ORG\/WIKI\/SAPPANWOOD Figure 8. Photomacrograph of Rubia tinctorum dried roots for prepared dyes and examples of madder dyes on paper. Figure 9. Sappanwood (Caesalpinia sappan). Photomacrograph of sappanwood and of paper dyed with sappanwood extract. KEITH LAWRENCE, MFA, BOSTON, HTTP:\/\/CAMEO.MFA.ORG\/WIKI\/TURMERIC 24 Arnoldia 74\/3 ? February 2017 Figure 10. Turmeric (Curcuma longa). Photomacrograph of dried and cut rhizomes and an example of turmeric dye on paper. Kiyomitsu I, turmeric is used as a clear, intense yellow for the trim as a contrast to the bright red. In the print by Kiyonaga, turmeric is used as a strong yellow background color. This colorant is fairly lightfast and retains its color better than the flavonoids. Flavonoids--silver grass: kariyasu; Japanese pagoda tree: enju; Toringo crabapple: zumi Flavonoids occur in most dye plants and their yellow colorants were discovered from the earliest times. While many colorants in this group are not lightfast, their abundance has resulted in their wide use. In Asia, the primary flavonoid-containing plants were the luteolin containing grasses, such as Miscanthus tinctorius (silver grass: kariyasu), that were cut each fall, then dried, and kept until the next spring for extraction. Other common Japanese dyeing plants include Styphnolobium japonicum (syn. Sophora japonica; Japanese pagoda tree: enju, see figure 11) and Malus sieboldii (Toringo crabapple: zumi). Flavonoid-containing dyes tend to have numerous compounds. For fluorescence measurements, the emission positions were similar and tended to blend into a single elongated peak. This pattern tends to be weaker than the turmeric pattern and was often noted mixed in with the absorption pattern for the paper, thus making positive identification difficult. Additionally, it was difficult to make any consistent determination for the various types of flavonoid yellows. Since each contains similar compounds, but in different proportions, the excitation and emission maxima are similar, blending into an oblong mesa-type absorption area rather than a single peak. In this set of analyzed woodblock prints, flavonoids tended to be a popular early colorant that later gave way to the use of turmeric and orpiment. One possible reason for the shift in yellow colorants is the poor light stability of most flavonoid yellows. The prints shown in figures 1, 4, 5 and 6 show examples of yellow flavonoid printed colors. Figure 6 was included as an example to show that even though it is difficult to distinguish between various flavonoid colors, this print does show two visually different yellow-colored regions that both produced slightly different fluorescence spectra even though both corresponded to flavonoids. It is possible that this print contains two types of flavonoid yellows, such as Japanese pagoda tree and Toringo crabapple. It is also possible that the print was exposed to uneven levels of light and that the yellows at the top of the print have deteriorated differently than those at the bottom. KEITH LAWRENCE, MFA, BOSTON, HTTP:\/\/CAMEO.MFA.ORG\/WIKI\/PAGODA_TREE Plant Dye Identification 25 KEITH LAWRENCE, MFA, BOSTON, HTTP:\/\/CAMEO.MFA.ORG\/WIKI\/GAMBOGE Figure 11. Japanese pagoda tree (Styphnolobium japonicum) flower buds and photomacrograph of paper dyed with pagoda tree buds. Figure 12. Photomacrograph of Garcinia hanburyi (gamboge) resinous pieces from the Harvard Museum of Natural History and photomacrograph of paper dyed with gamboge. Gamboge: shio Gamboge is a golden yellow colorant that is extracted by tapping resin from various species of the evergreen Garcinia trees, most commonly G. hanburyi and G. morella native to southeast Asia and India (see figure 12). The trees must be at least ten years old before they are tapped. The resin is extracted by making spiral incisions in the bark, and by breaking off leaves and shoots and letting the milky yellow resinous gum drip out. The resulting latex is collected in hollow bamboo canes. After the resin is congealed, the bamboo is broken away and large rods of solidified resin remain. Gamboge is marketed as solid pieces or as a powder. Visually, under a hand-held UV light, the yellow regions containing this colorant appeared dark, as if absorbing the fluorescence. The lack 26 Arnoldia 74\/3 ? February 2017 of fluorescence was confirmed by the EEM fluorescence pattern produced for reference samples of gamboge. The pattern exhibited a complete absence of emission peaks; this also included an absence of the paper peaks indicating that the paper was covered with a blocking agent. Gamboge is the only known organic reference in the potential set of Japanese colorants that corresponds to this negative pattern. Thus, this is a unique material that visually appeared yellow, but without sampling, it could only be characterized by the absence of any measurable inorganic elements (e.g., Fe, As) by XRF along with the absence of any unique fluorescence pattern by EEM fluorescence. Gamboge was most often found in the handcolored, beni-e, prints and is illustrated in the corner of the warrior's robe (figure1), along with other non-analyzed points such as the kimono collars and the center frame of the umbrella. Dayflower: aigama; awobama Indigo: ai Indigoid dyes were used in Neolithic Europe, Pharaonic Egypt, and now in twenty-first century jeans (see figure 15). While the source plants provide slightly different hues, indigo REBEKAH D. WALLACE, UNIVERSITY OF GEORGIA, BUGWOOD.ORG KEITH LAWRENCE, MFA, BOSTON, HTTP:\/\/CAMEO.MFA.ORG\/WIKI\/FILE:DAYFLOWER_BLUE.JPG Though rarely used elsewhere, dayflower (Commelina communis) blue was commonly used in Japan. Dayflower is an annual plant native throughout much of eastern Asia that bears one-day-blooming flowers featuring two large blue upper petals. The anthocyanin-containing juice extracted from the flowers was used by illustrators and printers for blue and green colors. Cloth or paper was dipped into the juice and dried; once needed the cloth or paper was dipped into water to extract the blue colorant (see figure 13). The best analytical method for the identification of dayflower is fiber optic reflectance. An example of the difference in the FORS spectra for dayflower and indigo is shown in figure 14. As dayflower and indigo were the only two plant-based blues dyes used for woodblock prints, the FORS method could quickly and simply distinguish between the two materials. Though dayflower was sometimes used by itself for blue areas (see figures 3 and 6), its poor lightfastness and its sensitivity to water were possible reasons that it was most often found used for greens and purples. Figures 3, 4, 5, and 6 show examples of the green and purple tones. Figure 13. Dayflower (Commelina communis) plant photo and photomacrograph of water soluble dayflower blue dried on paper. Plant Dye Identification 27 FORS spectra 120 100 80 dayflower indigo 60 40 colorless soluble form. Once the colorant is extracted, it is either printed out or cast into cakes where the insoluble blue indigo precipitates as it reacts with oxygen from the air. Of the prints selected for this article, indigo was found in figures 4 and 5 in the bright grass green colors. For both prints, orpiment, an inorganic yellow, was mixed with indigo to obtain the vivid, somewhat lightfast color. Conclusion 20 995 910 825 740 655 570 485 400 0 Figure 14. Overlay graph showing the Fiber Optic Reflectance (FORS) spectra for dayflower (Commelina communis) on paper versus indigo (Indigofera tinctoria) on paper. KEITH LAWRENCE, MFA, BOSTON, HTTP:\/\/CAMEO.MFA.ORG\/WIKI\/INDIGO has been regarded as the color of kings. Indigo producing plants, such as Indigofera tinctoria (a tropical shrub or subshrub), contain colorless glycocides that can be converted to the blue colored indigo on exposure to oxygen. To produce the dye, the fresh leaves are macerated in hot water, after which an alkali is added (such as lime) to ensure the colorant remains in a The purpose of this paper is to provide specific information on the analysis and identification of natural colorants used in the production of Japanese woodblock prints. Three nondestructive analysis techniques were used so that no samples were removed from the prints. X-ray fluorescence (XRF) was used to determine the presence of any inorganic compounds, and fiber optic reflectance spectroscopy (FORS) was used to distinguish between indigo and dayflower in the blue, green, and purple regions. Additionally, methods were developed to successfully use a third technique, excitationemission matrix (EEM), or 3-D, fluorescence spectroscopy, for the characterization of the red and yellow plant-based colorants. Figure 15. Indigo (Indigofera tinctoria). Photomacrograph of powdered indigo dye and of paper dyed with indigo. 28 Arnoldia 74\/3 ? February 2017 The MFA collection of Japanese woodblock prints is an ideal venue for the use of three combined techniques for the identification of the colorants because: ?The palette used for woodblock prints is limited. ? The colorants and substrates for the print were prepared with consistent, often documented, methods that had minimal variation. ?The prints are flat and the size of the prints, even within their mats, is less than 1 square meter. ?The speed for all three types of analysis is fast and allows for easy analysis of multiple locations. ?The MFA has an extensive set of over 50,000 Japanese woodblock prints and illustrated books that allows for exten- sive surveys of the materials within each time period, style, publisher, and artist. ?The knowledge obtained from the colorant identification will promote the understanding of the light stability for each print, and thus help preserve its vibrancy. Acknowledgements The authors are grateful to Kathryn Richardson (The Arnold Arboretum) and Allaire Diamond (Williston, Vermont) for supplying known reference samples for analysis. Additionally, we commend Arianna McQuillen and Kaeley Ferguson for conscientiously analyzing the set of Japanese prints by EEM fluorescence and FORS. Michele Derrick is the Schorr Family Associate Research Scientist, Joan Wright is the Bettina Burr Conservator in Asian Conservation, and Richard Newman is the Head of the Scientific Research Division, all at the Museum of Fine Arts, Boston. 36673667 U.S. POSTAL SERVICESTATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION(Required by 39 U.S.C. 3685) 1. Publication Title: Arnoldia. 2. Publication No: 0004?2633. 3. Filing Date: November 2, 2016. 4. Issue Frequency: Quarterly. 5. No. of Issues Published Annually: 4. 6. Annual Subscription Price: $20.00 domestic; $25.00 foreign. 7. Complete Mailing Address of Known Office of Publication: Arnold Arboretum, 125 Arborway, Boston, Suffolk County, MA 02130?3500. 8. 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Average No. Copies Each Issue During Preceding 12 Months: 80%. Actual No. Copies of Single Issue Published Nearest to Filing Date: 77%. I certify that all information furnished on this form is true and complete. Nancy Rose, Editor. "},{"has_event_date":0,"type":"arnoldia","title":"A Tour of the Oaks of the Arboretum des Pouyouleix","article_sequence":3,"start_page":29,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25614","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14ea728.jpg","volume":74,"issue_number":3,"year":2017,"series":null,"season":null,"authors":"Chasse, Beatrice","article_content":"A Tour of the Oaks of the Arboretum des Pouyouleix B?atrice Chass? O for a worldwide total of 430. Visitors to the Arboretum des Pouyouleix are always surprised to learn that Mexico, with 150 (90 of which are endemic), is the country with the greatest number of species. The diversity in number of species is paralleled by the morphological diversity of the EIKE JABLONSKI aks (Quercus) occupy every ecological niche in the Northern Hemisphere. The natural distribution of the genus only extends into the Southern Hemisphere in Indonesia where a few species can be found. In Europe there are 38 species, in Asia 156, while North and South America together harbor 236, Oak (Quercus) acorns are diverse in size and form. Clockwise from upper left: Q. insignis (Mexico); Q. chrysolepis (USA); Q. hypoleucoides (USA); Q. dolicholepis (Asia); Q. monimotricha (Asia); Q. macrolepis (Europe) ALL PHOTOS BY THE AUTHOR UNLESS OTHERWISE INDICATED 30 Arnoldia 74\/3 ? February 2017 Leaves of the Mexican oak species Q. urbanii are bright crimson when newly emerged. The handsome form and foliage of loquat oak (Q. rysophylla) has made it a favorite of plant connoisseurs. Quercus viminea, native to Mexico, has elegant, glossy foliage. Quercus cornelius-mulleri is a scrub oak native to California and Baja California. It was named for noted botanist and ecologist Cornelius H. Muller (1909?1997). Oaks of the Arboretum des Pouyouleix 31 leaves, as reflected by some of their common names: maple-leafed oak, loquat oak, chestnut oak, bamboo oak, holly oak, willow oak, myrtle-leafed oak, laurel oak, and so on. The diversity of acorn morphology is as surprising and wonderful as the diversity of leaf morphology and habit. Since the oak collection of the Arboretum des Pouyouleix is planted geographically, I'd like to introduce Arnoldia readers to some of the most interesting oaks here by taking a tour through the collection and providing some details and personal memories of collecting trips around the world. MEXICO Though I find it impossible to decide which is my single favorite oak, without a doubt my favorite oaks come from Mexico. Beyond their extraordinary diversity and beauty, my fondness results from the facts that, one, many of my best oak adventures with both the plants and the people interested in those plants are linked to that country, and two, that all of the Mexican oaks, with a few exceptions, grow so well here at the Arboretum des Pouyouleix. To be sure, there are beautiful and interesting oaks all over the world but for me nothing quite so extraordinary as a young leaf of Q. urbanii, as unbelievable as the acorn of Q. insignis, as elegant as Q. viminea, as majestic as Q. rysophylla--the list of superlatives is endless. Although it might be an exaggeration to say that Q. rysophylla, loquat oak, is everybody's favorite tree, it does have an impressive list of admirers. It was selected as \"Tree of the Year\" in 2015 by the International Dendrology Society (IDS), and botanist Allen Coombes, writing for the IDS Yearbook, described the young coppery leaves as quite unlike anything he had ever seen before when he first saw the tree in 1980 (Coombes 2016). John Grimshaw, in New Trees: Recent Introductions to Cultivation (Grimshaw and Bayton 2009), wrote, \"Of all the trees in this book, Quercus rysophylla is the one that has made the greatest impression on me, wherever it has been seen, and if only one `new tree' were to be grown, this should perhaps be it.\" In 1978, Lynn Lowery, horticulturist and plant collector from Texas, regarded this species as a VIP (very important plant) (Creech 2016) and John Fairey, another noted plant collector and nurseryman from Texas, said \"If I had to have one oak, it would be rysophylla.\" (Raver 2012). Our first Q. rysophylla was grown from acorns collected in Chipinque Park in Nuevo L?on, Mexico, and planted here in 2004, measuring 23 centimeters (9 inches) tall. Today it measures nearly 9 meters (30 feet). The lovely dark green, thick and shiny, nearly sessile leaves are densely clustered and, when young, vary in color from bronze to red. We have planted six other trees of this species, which have grown even faster. \"9,490 Kilometers Across Mexico\" (Chass? 2011), an account of my second trip to Mexico, in 2010, could just as well have been titled \"In the Footsteps of Cornelius H. Muller.\" Perhaps Quercus mulleri and Q. cornelius-mulleri are the most immediately visible traces in the oak world of this incredible botanist who was still actively involved, in his mid-eighties, in preparing the Quercus section of the Flora of North America with botanist Kevin Nixon. Muller was indefatigable in the field and a large part of my itinerary in 2010 was based on the detailed location descriptions of the oak discoveries he made during his adventures in northern Mexico. These included Q. ? basaseachicensis near the Basaseachic Falls in Chihuahua, Q. flocculenta halfway up the Cerro Potos? in Nuevo L?on, Q. edwardsiae in Chipinque, Nuevo L?on, but above all, in Chihuahua, Q. deliquescens, which provides a story that started in the town of Delicias (\"delight\" in Spanish) and ended in Milagro (\"a miracle\"). Muller wrote, \"Recent heavy concentration of collecting efforts in the Chihuahuan Desert region of Mexico ... have yielded much new information on the flora and its distribution. Among the novelties is a striking species of Quercus here described as new\" (Muller 1979). The text continues with a precise description of how to approach the location (most useful even when one has GPS coordinates) and so off we went, leaving Delicias for the town of Julimes to get to the Sierra del Roque \"... as approached from Minas Las Playas via Rancho El Saucito.\" After several hours of very difficult and slow driving through the Chihuahuan Desert, The Arboretum des Pouyouleix, National Oak Collection (France) How does one become an oak nut? Initially, my motivation was simply to create a botanic garden. But while my companion and I drove around southwestern France in search of suitable land, we kept noticing a majestic tree that dominated the landscape--it was Quercus robur. Though perhaps best known as pedunculate or English oak, it is such a common tree in northern Europe that it has dozens of common names in many languages. This inspired us to make oaks the focus of our future garden. The Arboretum des Pouyouleix is located in the north of the Aquitaine region of France, in the department of the Dordogne, roughly 150 kilometers (93 miles) north of the city of Bordeaux and 200 kilometers (124 miles) from the western coast of France. The topography is quite variable, which provides planting sites with differences in soil composition and structure, drainage, exposure (to both wind and sun), and temperatures. Although we are theoretically in USDA Hardiness Zone 8b (average annual minimum temperature -9.4 to -6.7?C [15 to 20?F]), we rarely experience winter temperatures lower than -4?C (24.8?F), and summer highs are quite often in the mid 20s to 30?C (77 to 86?F). The average annual rainfall (for the past 11 years) is 917 millimeters (36.1 inches). We decided to create the collection with plants raised from seeds, and preferably from wild-collected seeds in order to reduce (though not entirely eliminate) the possibility of hybridization. A propitious encounter in 2005 with several European oak enthusiasts from the International Oak Society paved the way for a series of collecting trips that have taken me across North America, Mexico, Vietnam, and Taiwan, collecting dozens of species including several new to cultivation in Europe. The Arboretum des Pouyouleix now holds a little over 300 Quercus taxa (including 38 species on the IUCN Red List) and is certified as a National Collection for the genus in France. In addition, the Arboretum has about 600 taxa in a variety of other genera. The table below shows the growth rate of seven Quercus species at the Arboretum des Pouyouleix. DATE PLANTED AND HEIGHT (m) NAME HEIGHT (m) 2015 AVERAGE ANNUAL GROWTH Q. imbricaria1 12\/20032.00 11.00 0.75 Q. saltillensis2 11\/20110.09 3.50 0.90 Q. rysophylla2 11\/2004 8.00 0.70 11\/20040.87 6.00 0.50 Q. dentata3 0.23 Q. mexicana2 06\/20120.40 5.00 1.50 myrtifolia1 03\/20080.11 3.50 0.50 Q. hintoniorum2 03\/20070.10 4.50 0.60 Q. Native to: 1North America; 2Mexico; 3Asia Oaks of the Arboretum des Pouyouleix 33 The Arboretum des Pouyouleix has varied topography that provides ideal sites for many oak species. and, according to Mr. Muller's coordinates, just a hop, skip, and a jump from Q. deliquescens, we found ourselves confronted with a difficult choice: there before us, for as far as the eye could see in either direction, stretched a very tall barbed-wire fence. To go or not to go over the fence? What would you have done? Driving back to civilization, we realized that we had started off without thinking to bring any food with us, although we did fortunately have enough to drink. So it was a miracle indeed, that the first town we came to, after several more hours of driving, had a small restaurant named ... El Milagro! But, truth to tell, the real miracle was that we were luckier than Mr. Muller who ends his description of the species with \"... acorns unknown.\" This species is considered vulnerable by the International Union for Conservation of Nature (IUCN). The area devoted to Mexico is the largest part of the Arboretum des Pouyouleix and comprises Quercus deliquescens is a rare oak species native to Chihuahua, Mexico. 34 Arnoldia 74\/3 ? February 2017 India) are the two most common oaks from this group found in collections in Europe, the former having been introduced to cultivation in 1854 and the latter in 1804. We have several trees of both of these species that grow well. Q. myrsinifolia makes a prettier tree here, whereas Q. glauca tends to be bushy. This group of oaks is not very well represented in American arboreta. Of the 20 gardens in the Plant Collections Network (PCN) Quercus Multisite Collection, the University of Washington Botanic Garden has four, Bartlett Tree Research Lab (North Carolina) has five, and the Scott Arboretum (Pennsylvania), the University of California?Davis Abundant male flowers are seen on the Mexican species Q. hintoniorum. Arboretum, and the Morris Arboretum the greatest number of taxa. Many delightful (Pennsylvania) each has one. Though not part oaks rare in cultivation can be found here: Q. of the Multisite Collection, the Aiken Citywide macvaughii; Q. miquihuanensis, an endangered Arboretum (South Carolina) also has four. species; Q. hintoniorum, listed as vulnerable; Q. My two favorite ring-cupped oaks growing crassifolia; Q. furfuraceae, listed as likely endanhere are Q. salicina (Japan, maybe Taiwan) gered; and Q. liebmannii, to name but a few. and Q. gilva (Japan, Southern China, Taiwan, Vietnam). Quercus salicina is just a perfect, ASIA graceful tree. The leaves, evergreen (as with all Current phylogenetic understanding of the Cyclobalanopsis), are elegantly elongated with genus Quercus is that it is composed of eight a twist at the end. It will produce new leaves at lineages or groups. The group known as the various times from spring until early autumn, ring-cupped oaks (section Cyclobalanopsis) is coloring the tree to different degrees in a beautiful deep burgundy red that fades to pink and only found in Asia. Not all of the oaks that grow finally to green. Of all our evergreen species it in Asia belong to this group--some of them is one of the few that suffered absolutely no belong to the white oak (section Quercus) lineage, which is ubiquitous throughout the natudamage during a horrific 15-day cold spell in ral distribution. Hands down, the ring-cupped February 2012 with temperatures at night dropping to -18?C (-0.4?F), and daytime temperaoaks would win first prize in a contest for the tures never above -8?C (17.6?F). Quercus gilva most un-oak-like plants (at least for Western is one of the more easily recognizable Cycloeyes), just as they would also win the contest balanopsis in part because its leaves are characfor the group whose members are the hardest to teristically widest in the middle but especially distinguish from one another. Come to think of because the new leaves and shoots are distinctly it, the seeds of at least two species, Q. macrocalyx (China, Southeast Asia) and Q. pachyloma yellow with a soft tomentum, giving it a unique (Southern China, Taiwan), would probably also ornamental quality win first prize in an acorn beauty contest. These The Arboretum's Asia section also has many two species, collected in Vietnam and Taiwan, Asian oaks that are in sections other than respectively, are still in the nursery, perhaps to Cyclobalanopsis: Q. dentata, with its huge, be planted this year. leathery leaves and sculptured bark; Q. spinosa, Quercus myrsinifolia (China, Japan, Southa very rare oak in cultivation; and Q. semecareast Asia) and Q. glauca (China, Japan, Southpifolia, which holds, along with Q. guyavifolia east Asia, Afghanistan, Bhutan, Nepal, Sikkim, and Q. monimotricha, the oak record for high- CHARLES SNYERS D'ATTENHOVEN Oaks of the Arboretum des Pouyouleix 35 Asian oak species Q. macrocalyx (left, photographed in Vietnam) and Q. pachyloma (right, photographed in Taiwan) have fabulous acorns. Asian evergreen oak species Q. salicina (left) and Q. gilva (right) have handsome foliage. altitude living (up to 4,000 meters [13,123 feet] for the first two and 4,600 meters [15,092 feet] for the latter). Our expedition to Vietnam in 2013, though not entirely satisfactory in terms of the number of oaks found, was most interesting in what it revealed about the presence of certain oaks hitherto unreported in northern Vietnam (Chass? 2014). Much remains to be learned about the status of the oaks of Vietnam, indeed, about the forest communities in general, since during the second half of the twentieth century war, forest fires, slash and burn agriculture, encroachment for industrial purposes, and other forms of anthropic pressure have reduced forestland in Vietnam from 43% of the country's surface area in 1940 to 17% by the late 1970s (Bien 2001). Daimyo oak (Quercus dentata), native to Japan, Korea, and China, bears enormous leaves. 36 Arnoldia 74\/3 ? February 2017 CHARLES SNYERS D'ATTENHOVEN EIKE JABLONSKI EUROPE Oak species diversity in Europe is not very high but there is interesting morphological diversity within the species present, and a few of them can indeed become most spectacular trees with truly impressive lifespans, especially Q. robur, pedunculate or English oak. This species also has the honor of being the first oak to have had its genome entirely sequenced (Plomion et al. 2015), a milestone for research into the evolutionary history of the genus. From an aesthetic point of view, the problem with quite a few European oaks is that they are moderately to severely affected by many diseases (powdery mildew, rusts, etc.) that, although not lethal, make the trees rather unattractive fairly quickly after the appearance of new foliage in spring. Quercus alnifolia, endemic to Cyprus, is one of my favorites with its golden yellow to orange tomentum on the underside of the round and shiny evergreen leaves and its fabulously elegant acorns. It is a large shrub or small tree (6 to 9 meters [20 to 30 feet]) and as such makes a wonderful addition to any small or medium-sized garden. Q. frainetto (Balkans, Bulgaria, Greece, Hungary, Italy, Romania, and Turkey) is another very special European oak, the deep sinuses of the leaves giving a delicate lacelike aspect to the silhouette. This species is also less prone to the above-mentioned afflictions. But my vote for the prettiest of all European oaks would be Q. macrolepis. Found across southeastern Europe from the Balkans to the Aegean Sea, Italy, and Turkey, it can be a shrubby tree of 5 meters (16 feet) or attain grand heights of 25 meters (82 feet) or more. The very striking silvery, grayish white color of the new leaves makes it a true eye-catcher on sunny spring days. And then of course, there is the acorn: one of the most fabulous of the genus (see page 29). I think that part of my enchantment with this species comes not only from its beauty but also from the importance of these acorns in human history, both in the leather-tanning industry for more than four hundred years (Mayer Maroulis 2014) and as a food source for probably much longer than that (Chass? 2016). Interesting European oak species include (top to bottom) Q. alnifolia (photographed in Cyprus), Q. frainetto (photographed at Wisley, United Kingdom), and Q. macrolepis. Oaks of the Arboretum des Pouyouleix 37 NORTH AMERICA We move now into the North American section. A dry, rocky, poor-soil area in this part of the Arboretum has proven to be an ideal place to plant many oaks from the southern (both east and west) United States. Generally, these oaks are accustomed to some level of environmental stress such as little rain, few nutrients, or harsh sun exposure. They are healthy plants here in France, many of them fruiting after only a few years, but tend to be slow growers. Four specimens of my favorite one, Q. palmeri, raised from seed collected in 2007 in Riverside County, California, were planted here in 2008, each measuring about 8 centimeters tall. Today they are all about 1 New foliage and flower buds of Q. palmeri, a shrubby oak native to meter tall (trees of this shrubby species California, Arizona, New Mexico, and Baja California. are generally not more than 3 meters tall oak (Q. nigra), swamp white oak (Q. bicolor), at maturity). The emblematic southern live oak white oak (Q. alba), black oak (Q. velutina), bur (Q. virginiana), one of the most magnificent oak (Q. macrocarpa), pin oak (Q. palustris), and oaks of the southern United States, with its many, many others are thriving here, reaching wide-spreading branches that are often dripping heights (for the fastest growers) of more than with Spanish moss in its natural habitat from 15 meters (49 feet) since 2003. This area of the Texas to Florida and northward to Virginia, Arboretum is the best seat in the house come does very well here, as do Q. chapmanii, which autumn as this mix of trees produces a vertigiis also from the southeast, Q. toumeyi from nous scale of color from yellow to orange to Arizona, and Q. engelmannii from California pink to red, set off by the surrounding chestnut(listed as vulnerable on the IUCN Red List). Nearby is an area with deep, rich soil that oak woodland that gives a magnificent backwe call \"la Grande Prairie.\" It was the first part drop of yellow and orangish brown. of the Arboretum to be planted, on December It is in this part of the Arboretum that I am 7, 2003, with the help of many neighbors and occasionally struck by the awesome temporal friends. We planted nearly 300 trees, most of dimension of what it means to plant trees. In them here. Under a bright blue sunny sky we just thirteen years, these trees, destined to live toiled away and, when we were finished, it several hundred years, have created a world of began to rain. Though I am not at all a superstitheir own but of which I am a part. And though, tious person, one has to admit that this was a regrettably, I will not live for several hundred good omen! In this part of the Arboretum can years, it is as though my trees have created a be found most of the common North American bridge through time for me. This exceptional oaks. I use the word \"common\" in the sense experience was magnified a hundred-fold when that they have been in cultivation for a long I visited the Arnold Arboretum in October 2015 time--some for more than two centuries--and because, of course, many of the trees there (oaks can be easily obtained through the nursery and other) have been planted for decades, some trade, but they are nonetheless extraordinary for more than a century. They are thus at once trees. Northern pin oak (Q. ellipsoidalis), a bridge through time past and time future. For scarlet oak (Q. coccinea), Shumard oak (Q. shuthose of us who plant trees, the Arnold is truly mardii), willow oak (Q. phellos), shingle oak (Q. a unique voyage, and without a doubt the most imbricaria), northern red oak (Q. rubra), water magnificent arboretum I have ever visited. 38 Arnoldia 74\/3 ? February 2017 Many of the North American oaks display bright autumn leaf color. From right to left, going down the hill, Q. palustris, Q. coccinea, and Q. muehlenbergii. On either side of \"la Grande Prairie,\" moderately steep slopes with rather poor soil have provided well-drained, sheltered areas for the planting of more fragile species from North America. I have come to the conclusion that, although cold is obviously a limiting factor to plant survival, heavy, rich soil and too much water in the fall and winter are equally serious handicaps for a great many oaks. On these slopes, many oaks that probably shouldn't enjoy being here because of the cold are very happy indeed: Q. myrtifolia (fruiting this year) and Q. inopina from Florida, and Q. invaginata and Q. insignis from Mexico, to name but a few. CONCLUSION As we turn to walk back up \"la Grande Prairie\" I should just point out a fine specimen of Q. tomentella, an oak endemic to the Channel Islands (off the coast of southern California), considered to be vulnerable by the IUCN. Near the house and other buildings, at the top of \"la Grande Prairie,\" many oaks, irrespective of geographic origin but that share the characteristic of being rather short and liking fairly poor soil, have been planted, including Q. vacciniifolia from California, Q. pumila from Georgia, Q. minima from South Carolina, and Q. guyavifolia and Q. monimotricha from China. Many, many seeds have been--and are being--sown here at the Arboretum des Pouyouleix. The seeds collected during the trips I've mentioned in this article, and plants raised from them, have been shared with different gardens and arboreta in France and around the world (Argentina, Belgium, China, the Czech Republic, Germany, the Netherlands, Spain, Taiwan, the United Kingdom, the United Oaks of the Arboretum des Pouyouleix 39 States of America, and Uruguay). A few years ago, there were almost exclusively Mexican and North American species in our nursery: these days, almost only Asian species, with many wonderfully exciting plants from Vietnam, China, and Taiwan, including some that have yet to be identified. In these times of ecological crisis, I should like to end this little journey with a sincere homage to plant collectors and plant propagators past and present. Identifying fragile zones and endangered species is surely a useful exercise, but is it not necessary, if we want to save those species, to have knowledge about their cultivation? In Europe, very nearly the only arboreta that are growing Runner oak (Q. pumila) is a shrubby, spreading oak native to the Coastal many endangered oak taxa, or, less Plain of the southeastern United States. dramatically, the more recent introCoombes, A. J. 2016. Tree of the year. Q. rysophylla ductions, are the private ones. These collecWeath. International Dendrology Society tions are a valuable resource for conservation Yearbook 2015. pp. 22?52. efforts and for building awareness about the beauty and the diversity of our planet. Creech, D. 2016. Quercus rysophylla ? One Mexico Acknowledgements I would like to thank Kyle Port for soliciting this article, Cindy Newlander for providing the information on the PCN Quercus Multisite Collection, and the International Oak Society. Thanks in great part to this group-- originally founded to facilitate seed exchange--the world of oak enthusiasts is one of sharing and camaraderie and it is safe to say that without the Society, oak collections around the world, including that of the Arboretum des Pouyouleix, would not be what they are today. Works cited Bien, N. N. 2001. Forest Management Systems in the Uplands of Vietnam: social, economic and environmental perspectives. Report prepared by the Economy and Environment Program for Southeast Asia. Singapore. Chass?, B. 2011. 9,490 Kilometers Across Mexico. International Oaks 22: 70?88. Chass?, B. 2014. In Search of Vietnam's Elusive Oaks. Chass?-Colin-Snyers Vietnam 2014 (CCSV13). International Oaks 25: 135?161. Chass?, B. 2016. Eating Acorns: What Story Do the Distant, Far and Near Past Tell Us, and Why? International Oaks 27: 107?135. Oak that's Texas Tough. https:\/\/dcreechsite. wordpress.com. Grimshaw, J., and R. Bayton. 2009. New Trees: Recent Introductions to Cultivation. Richmond: Kew Publishing, Royal Botanic Gardens, Kew. Mayer Maroulis, M. 2014. Creating Sustainable Income From the Ancient Oak Forest on Kea Island, Greece. International Oaks 25: 13?22. Muller, C.H. 1979. Quercus deliquescens, A New Species from Chihuahua, Mexico. Phytologia 42: 289?291. Plomion, C., et al. 2015. Decoding the oak genome: public release of sequence data, assembly, annotation and publication strategies. Molecular Ecology Resources, doi: 10.1111\/1755-0998.12425 Raver, A. 2012. Growing From His Mistakes. A Texas Gardener Looks to Mexico for Inspiration. h t t p : \/ \/ w w w. n y t i m e s . c o m \/ 2 0 1 2 \/ 0 4 \/ 1 9 . Accessed on 15 September 2016. B?atrice Chass? is co-founder of the Arboretum des Pouyouleix, past President of the International Oak Society, and Editor of International Oaks, the journal of the IOS, since 2013. "},{"has_event_date":0,"type":"arnoldia","title":"One of a Kind: Pinus monophylla","article_sequence":4,"start_page":40,"end_page":40,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25615","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14ea76d.jpg","volume":74,"issue_number":3,"year":2017,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"One of a Kind: Pinus monophylla Nancy Rose I 've led many plant identification classes and walks in my career as a horticulturist. When it comes to pines (Pinus), I've taught that pines always carry multiple needles grouped in fascicles (bundles), which readily differentiates them from spruces (Picea) and firs (Abies) (the other common \"tall, pointy evergreens\"), which both bear single needles. To then identify individual pine species, the first step is to see if the bundles hold two, three, or five needles. That's still good advice about 99 percent of the time, but when I came to the Arnold Arboretum I discovered a notable exception to those rules: Pinus monophylla, the single-leaf pine. Pinus monophylla is a member of the pine family (Pinaceae) and is one of 114 species in the genus Pinus. It is part of subsection Cembroides, a group of pine species native to the western United States and Mexico commonly known as pinyons, or pi?ons (P. monophylla is also known as single-leaf pinyon). The pinyons have thin-shelled, edible seeds; P. monophylla and P. edulis, Colorado or two-needle pinyon, have especially large (1\/2 inch [1.3 centimeters] long), high-fat-content seeds that have long been harvested and used as an important food source by indigenous peoples. Single-leaf pine grows about 15 to 30 feet (4.6 to 9.1 meters) tall and has a pyramidal form when young, becoming more irregular and spreading with age. As the common name indicates, this pine bears single needles on its stems, the only pine species to do so. Individual needles are thick, sharp-tipped, and bluish green with silvery stomatal lines (see inset photo on opposite page). Abundant small staminate (male) cones release pollen and the round female cones mature to about 1.5 to 2.5 inches long in two years. Pairs of large seeds are held in depressions on individual cone scales; seeds are readily eaten by many birds and other wildlife species, and animal seedcaching, especially by pinyon jays (Gymnorhinus cyanocephalus), is the primary means of seed dispersal for the species. Pinus monophylla grows in a semi-arid native range that runs from northern Baja California to southern and eastern California, Nevada, the southeastern corner of Idaho, western Utah, and parts of Arizona and New Mexico. It is cold hardy enough (USDA Zone 6, average annual minimum temperature -10 to 0?F [-23.3 to -17.8?C]) for Boston, but our much wetter climate may be part of the reason this pine has been difficult to grow at the Arboretum. We have tried a number of P. monophylla accessions through the years, the first one in 1908, but we currently have no living specimens in the collections. The last one was accession 400-88-B, which was a repropagation (by grafting) of a 1964 accession (287-64), which came from seeds wild collected in Nevada. Accession 400-88-B was a handsome, healthylooking specimen when I photographed it growing in the dwarf conifer terrace at the west end of the Leventritt Shrub and Vine Garden in May 2009 (opposite page). Unfortunately, just a few years later it went into severe decline and was removed; signs of root rot were noted on its removal. In its native range, single-leaf pine typically grows on very well-drained, gravelly slopes. The much greater annual rainfall and moister soils at the Arboretum may well have contributed to the demise of this and other specimens. This unique pine species will certainly return to the Arboretum soon. Pinus monophylla is one of the approximately 400 taxa targeted for acquisition in the ongoing Campaign for the Living Collections (see the complete list in Arnoldia 73\/3). We already have three seed accessions collected last year in Utah that are currently undergoing stratification in the Dana Greenhouses. And when we do have young single-leaf pines ready to move to the grounds, special care will be taken to place them in a site where, ideally, they will thrive for many years. Nancy Rose is the editor of Arnoldia. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23453","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14ea36e.jpg","title":"2017-74-3","volume":74,"issue_number":3,"year":2017,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Rooted in the Collections","article_sequence":1,"start_page":2,"end_page":14,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25612","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170896b.jpg","volume":74,"issue_number":2,"year":2016,"series":null,"season":null,"authors":"Gapinski, Andrew","article_content":"CAMPAIGN FOR THE LIVING COLLECTIONS Rooted in the Collections Andrew Gapinski We are now one year into the Campaign for the Living Collections, a ten-year initiative that will expand and refine the Arboretum's historically and scientifically valuable plant collections. Additional collecting trips this summer and fall have brought in new seed and plant accessions that will go through the Arboretum's propagation facilities, as described in the last issue of Arnoldia. In this issue, Manager of Horticulture Andrew Gapinski completes the Campaign article series by describing the process involved in moving plants to permanent locations on the grounds and the ongoing challenges of keeping the collections healthy and growing. F rom its origin to the Arnold Arboretum's propagation facilities, much time and many resources have been invested in the planning, acquisition, and production of an accession in preparation for its ultimate installation in the permanent collections. Successful establishment of new accessions and care of the Arboretum's 15,000 existing specimens takes a dedicated team of highly skilled horticulture professionals who are involved in aspects from site selection and planting to aesthetic and corrective pruning, soil health management, and attention to various plant stressors as part of the Arboretum's holistic Plant Health Care Program. Across our 281-acre landscape, we are preparing the grounds for a surge of new material as part of the Campaign for the Living Collections (Friedman et al. 2016)--an initiative to acquire and cultivate 400 target taxa over the next ten years. The Arboretum's historic collections scheme is based on the Bentham and Hooker system of plant taxonomy, devised in the late 1800s, with species grouped by genus in an evolutionary progression starting with the earliest of flowering plants, e.g., Magnolia, placed at the Arboretum's main gate. While taxonomic systems differ today, the Bentham and Hooker blueprint for incorporating new material into the permanent collections is generally still followed. Continued and expanded attention is also placed on utilization of the unique microenvironments, with their variable factors such as temperature, moisture, light, and soil type, that can be exploited for the successful cultivation of particular species. The Explorers Garden, nestled on the south side of Bussey Hill, represents one such area, long known as the spot for evaluating marginally hardy species not typically grown successfully in New England. New landscapes continue to be added, including the Leventritt Shrub and Vine Garden, dedicated in 2002, which arose out of a need for a space to feature shrubs and vines requiring full sun, and the landscape surrounding the Weld Hill Research Building, completed in 2011, which provides an opportunity for development of a new plant collection at the hub of the Arboretum's research programs. Whether sited in a particular location for taxonomic, thematic, aesthetic, or practical cultivation purposes, the placement of each new specimen into our historic landscape is part of The Explorers Garden's protected microclimate makes it an ideal site for trialing new accessions of unknown cold hardiness. This view shows (foreground to background) Rhododendron yedoense var. poukhanense, Chionanthus retusus, and Fothergilla hybrids. Photo by Richard Schulhof. CONOR M. GUIDARELLI 4 Arnoldia 74\/2 ? October 2016 Manager of Horticulture Andrew Gapinski, Manager of Plant ProductionTiffany Enzenbacher, and Manager of Plant Records Kyle Port compare Pinus cembra (accession 237-2001) siblings for overall health, vigor, form, and root development, assigning a ranking based on their condition. This information is used to determine whether each specimen is ready to be transplanted to the collections. a well-thought-out decision making process, executed with sound horticultural practice. Measure Twice, Cut Once After being cultivated in the Arboretum's Dana Greenhouses and surrounding nurseries for about three to seven years, the process of determining which specimens are ready to find their place in the permanent collections starts in August of each year. A review by greenhouse staff of all accessions in the production facilities is undertaken and recommendations are made as to whether an individual plant is large and healthy enough for installation. With this information in hand, the Managers of Plant Production and Horticulture and the Curator of Living Collections visit each specimen for a final determination. Ideally, multiple individuals within an accession have been successfully grown to ensure the best chance of that lineage surviving the production cycle and many years in the collections. A comparison between these siblings for overall health, vigor, form, and root development is made, and individuals are assigned a ranking based on their overall condition. At this point, it is also determined whether spring or fall transplanting is most appropriate for the species under review. For example, many oaks (Quercus spp.), beeches (Fagus spp.), and hornbeams (Carpinus spp.) can fare poorly when transplanted in autumn, while other plants, including many conifers, acclimate just fine. For the past several seasons, fall planting has been limited or deferred altogether because of prolonged summer droughts that have persisted well into autumn. When conditions are favorable, getting a jump on the transplanting list in fall helps with the work load of the busy spring season. Nevertheless, ignate nodes where new accessions within a the vast majority of transplants occur in the genus can be sited together outside of the core spring when warming days, cool nights, and collection. For example, we have been clustering individual Carpinus specimens at a few abundant precipitation create favorable rooting nodes on Peters Hill. conditions. Depending on how many sibling The Arboretum is divided into 71 horticulindividuals are needed for the permanent collections (typically three or four), surplus plants tural zones, each of which is assigned to one of may then be offered up to other botanical instiseven horticulturists responsible for the daily tutions. The sharing of specimens at this stage care of the collections within. Continuing of the process offers yet another opportunity into fall, field selection of the specific planting for material to be \"backed-up\" elsewhere in the location for each specimen involves the Manager of Horticulture, Curator of Living Collecevent of loss at the Arboretum. tions, and the horticulturist assigned to that With the list of graduates in hand, the process of finding planting locations begins. After zone. Each planting location is marked with nearly 150 years of collections development, a wooden stake and is labeled with the taxon finding locations for the approximately 250 and accession number to be planted. With the annual plant additions to the permanent colmajority of planting scheduled for the following lections is no easy task. On paper, specimens spring, horticulturists will follow up before the are first loosely assigned to particular areas of ground freezes and turn the soil in place to further mark the planting location, because stakes the Arboretum. A number of different parameters are considered, including the species' can easily be lost over winter. This step also taxonomic group (family, genus, etc.), known provides an opportunity for soil amendments winter hardiness, moisture requirements, colto be added as needed and makes for easier diglections value of that particular lineage, and ging in the spring as the freeze and thaw of the aesthetic and functional qualities of the species season loosens the turned soil. for use in various landscape projects across the The planting locations of the qualifiers (individual plants assigned identification letters A, grounds. Since our museum specimens are living organisms exposed to many environmental B, C, etc.) of accession 637-2010, a Yunnan redinfluences (drought, disease, winter storms, etc.), lineages of high value are sometimes duplicated across different parts of the Arboretum landscape as a means of internal backup. However, as a general rule, most of the plants within an individual accession are planted in the same collection area, with an occasional planting in an alternative section. Some designated areas, such as the Carpinus collection near Valley Road, are rather full of high-value trees and leave little room for development. When siting new accessions here, we may plant just one in this core collection area, and then plant the remaining two or three siblings together in another area. To avoid the look of random plants A flag and wooden stake mark a newly selected planting site for a mountain dotted through the landscape, we've maple (Acer spicatum, accession 270-2010-B) grown from seeds collected by recently begun to identify and des- Arboretum staff in New York's Adirondack Mountains. KYLE PORT Living Collections 5 Gaining Ground ANDREW GAPINSKI The Campaign for the Living Collections is under way and seeds from both near and far are sprouting in the Dana Greenhouses. As Manager of Horticulture, I can't help but feel a bit of anxious excitement as I await the challenge of growing new taxa from around the world. As stewards of the Campaign we face many questions including \"Where will we find the space and how can we prepare the grounds now to receive new material?\" As a horticulture team, we are viewing the grounds through an opportunistic lens--what is the value of each specimen to the collection, how can we better utilize the various environmental conditions we find across our landscape, how can we benefit from issues that affect the health of our collections, what areas need additional attention, and how can we gain \"new\" ground? Although not necessarily novel questions for collections managers to be considering, these concepts are at the forefront of our decision making across the landscape: Addition through subtraction Not every specimen in our collections holds the same value, and making tough decisions to deaccession and remove otherwise healthy plants is not easy, but for the building of any museum collection what is taken away can be just as important as what is added. We seek to utilize our limited resources, including space and staff time, in the most effective way to achieve the Arboretum mission. The decision-making process for the continued stewardship of every accession considers many factors, including total opportunity costs, and is ultimately guided by our Living Collections Policy (Arnold Arboretum 2016). Reclaiming areas of deferred maintenance Of top priority is the reclaiming of areas in which maintenance was deferred at some point. The Horticulture staff is undertaking an aggressive cleanup effort in collections areas that have been reabsorbed into adjacent natural lands and succumbed to invasive weeds. Over the past several decades, the southwest edge of the Hickory (Carya) Collection (seen here) has been slowly reclaimed by the adjacent Central Woodland--this area represents over an acre of valuable territory that will be available for collections expansion once it is cleared. ANDREW GAPINSKI The deaccessioning and removal in 2015 of these four cultivars of Norway maple (Acer platanoides), which had low collections value, freed up valuable acreage in the heart of the Maple (Acer) Collection for new high priority taxa. In the opening created by the removals, a purpleblow maple (Acer truncatum, 629-2010A), wild-collected in Shaanxi, China, during the September 2010 North America-China Plant Exploration Consortium (NACPEC) expedition, was planted. Our collections of trees, shrubs, and vines face continuous and ever-changing biotic and abiotic influences. When faced with events beyond our control, triaging the situation to prioritize and safeguard the most valuable holdings and finding opportunity in the loss is of utmost importance. ANDREW GAPINSKI The emerald ash borer (EAB), introduced to the United States from Asia via wood packaging material, has decimated ash (Fraxinus spp.) populations across central and eastern North America since its discovery in Michigan in 2002. Through the establishment of an early detection program in partnership with the Massachusetts Department of Resource Conservation, the Arboretum was the first detector of EAB in the City of Boston. A triage approach to evaluate the 146 accessioned ash trees in the permanent collections was undertaken. Fifty-one individuals were determined to be of low value, not warranting long-term preservation efforts. Of these, fifteen were immediately removed because of poor health, and the remaining thirty-six trees are being utilized for further EAB monitoring efforts. The remaining 95 high-value accessions are part of a preservation program that includes prophylactic treatments and clonal propagation efforts. Branch sections (including buds) from a select number of lineages were also sent to the USDA-ARS National Laboratory for Genetic Resources Preservation for potential long-term storage via cryopreservation. The photograph shows Arboretum Horticulturist Scott Grimshaw treating a high-value ash specimen. Exploring the potential of environmental niches Across the Arboretum, differences in environmental conditions from soil moisture to annual minimum temperature exist and are key considerations when selecting locations for species requiring particular niches. We are taking an aggressive approach to clean up these areas in preparation for harboring new accessions to come. The group project for the 2016 Isabella Welles Hunnewell Interns was to advance the development of the \"The Rockery,\" an area with exposed rock outcroppings along Valley Road (seen here, Arboretum Horticulturist Greg LaPlume removes excess soil from the site). Taking advantage of the natural geology of the site, the Arboretum seeks to develop an environment that supports species adapted to rocky mineral soils, with characteristically low nutrient and organic matter levels, such as those found in scree type habitats. Prickly-pear cactus (Opuntia humifusa), several ephedra (Ephedra) species, regal lily (Lilium regale), and a number of other species on the Campaign for the Living Collections list of desiderata are potential candidates for cultivation in such an environment. ANDREW GAPINSKI With loss comes opportunity: making lemonade out of lemons 8 Arnoldia 74\/2 ? October 2016 WILLIAM (NED) FRIEDMAN bud (Cercis glabra) collected by Michael Dosmann, Curator of Living Collections, on the September 2010 North America-China Plant Exploration Consortium (NACPEC) expedition to Shaanxi, China, serves as an example of the basic thought process for site selection. Having attempted to grow the species at the Arboretum several other times without success, the limited history of its cultivation here made planting decisions more difficult. Particularly in cases in which hardiness of the species is questionable, such as C. glabra, we use knowledge of the Arboretum's long studied and utilized microclimates to give us the best chance of success (Dosmann 2015). With the rolling topography, cold air drains down from the tops of the Arboretum's highest points including Bussey, Hemlock, and Peters Hills to the valleys below. In a typical year, these \"hot spots\" of the higher elevations experience minimum temperatures representative of a Zone 7 (average annual minimum temperatures 0 to 10?F [-17.8 to -12.2?C]), with Zone 6 (-10 to 0?F [-23.3 to -17.8?C]) conditions being most prevalent throughout the grounds. With six individuals of 637-2010 ready for the planting in the spring of 2015, what was the planting approach? Three were planted that spring: one (637-2010-A) in the microclimate of the Explorers Garden, located along Chinese Path on the south side of Bussey Hill, and the two others (B and C) among its relatives in the Legume Collection. The three remaining (D, E, and F) were held back in the greenhouses as reserves in case hardiness turned out to be an issue. The winter of 2015?2016 would turn out to be a true test of hardiness, with a season low of -14.5?F (-25.8?C; Zone 5) recorded in the Bradley Rosaceous Collection--the lowest temperature recorded at the Arboretum in 57 years. Spring 2016 came and observations were made; the Explorers Garden specimen leafed out fully with no dieback and the Legume Collection plants experienced only moderate branch dieback of 1 to 2 feet (30.5 to 61.0 centimeters). Success! With hardiness a non-issue, the three remaining plants were sited and planted in the landscape surrounding the Arboretum's Weld Hill Research Building for the species' ornamental value, its botanical and taxonomic interest, and the exploration story it brings to the newly developing Weld Hill landscape. The Weld Hill planting also is separated from its previously planted siblings by nearly a mile. That distance is a key part of the idea of internal back-up. Planting Season As spring approaches, we pay close attention to the thawing soils and moisture conditions and begin the transplanting process as soon as the timing is right. In preparation, planting lists and locations are reviewed, and a final walk-through of the nurseries is performed to document and adjust plans based on damage that may have occurred to plants over the winter. For example, following the record breaking snowfall--110.6 inches (280.9 centimeters) measured at Boston's Logan Airport--during the winter of 2014?2015, significant damage in the nurseries occurred as the snow melted and refroze during the spring thaw. Many young trees with low branches were pulled apart with the shifting snow and ice that covered them. Evaluations completed the Flowers of a Yunnan redbud specimen (Cercis glabra, 637-2010-D) that was planted in the Weld Hill Research building landscape. (continues on page 12) A Focus on Soil Health Jenna Zukswert, Living Collections Fellow In order for the nearly 400 taxa we intend to collect in the Campaign for the Living Collections to have a lasting legacy here at the Arboretum, we seek to provide specimens with the most favorable habitat possible. In preparation for the arrival of these new taxa, the horticulture department is investigating all aspects of plant health care and landscape management practices to ensure we are providing the highest of horticultural standards that will give these new accessions the greatest chance of success. Evaluation of the current conditions of our soils and the development and implementation of strategies to improve soil health across the Arboretum is a high priority. Awareness of the importance of soils, especially in relation to plant health, has been increasing; this was recently reflected in the United Nations-designated International Year of Soils in 2015. We as a global society are now more aware of how precious soil is, and also how often this resource is mismanaged. In addition to serving as a substantial terrestrial carbon sink, storing carbon that would otherwise be released into the atmosphere and further contribute to climate change (Dungait et al. 2012), soil provides important services that promote plant health, such as nutrient supply, water regulation, and physical support for roots (United States Department of Agriculture 2016). Therefore, proper stewardship of our soils will enhance the health of our valued Living Collections. The first step in caring for our soils is to fully understand their current state. In the late 1990s and early 2000s, an intensive program of soil assessment and management at the Arboretum identified highly acidic and nutrient deficient soils, which were at the time contributing to substantial tree decline in certain areas. In the years that followed, liming and nutrient applications were made, and some plant recovery was noted. Through staffing changes over the past decade, work on soils health waned, with only isolated issues addressed as plant decline was observed and investigated. With the onset of the Campaign, the Arboretum has renewed its commitment to understanding and managing the factors that affect soil health across our landscape. In 2015, with the help of that year's Isabella Welles Hunnewell Interns, Arboretum-wide soils testing that measured an assortment of nutrients and chemical characteristics was completed, as were analyses related to microbial processes. Initial recommendations for remediation were prescribed and executed. These soil testing data were recently mapped; the Geographic Information Systems (GIS) layer housing these data makes it possible to visualize all of the soil characteristics measured in the samples throughout the landscape. Having these data displayed spatially reveals landscape-wide patterns in soil health that we might otherwise overlook when referring only to a spreadsheet. We have also started to map the location of amendments that we apply to the soil; we hope that by continuing to measure and map soil characteristics and management efforts we can detect trends in soil health over time and determine the effectiveness of various management actions. We can use this new knowledge to adjust our practices and make more informed management decisions in the future. In addition to looking at these data ourselves, the formation is underway of a Soils Advisory Committee composed of soil science experts who can help us to further interpret these data and inform our management plans. Members of this committee contribute expertise from a wide range of topics in soil science and management, including bedrock geology, forest soil ecology, composting, and agricultural cover cropping. This committee will help connect us to academic research at Harvard University and beyond, enabling us to consider new ideas and to address with evidence the questions we have in our attempts to provide the best substrate for our expanding collections. N LEGEND Soil pH 4.07 - 4.28 4.28 - 4.49 4.49 - 4.71 4.71 - 4.92 4.92 - 5.13 5.13 - 5.34 5.34 - 5.55 5.55 - 5.77 5.77 - 5.98 5.98 - 6.19 JMZ, 8\/12\/2016 The Arnold Arboretum of Harvard University JMZ, 8\/12\/2016, THE ARNOLD ARBORETUM OF HARVARD UNIVERSITY 1 inch = 926 feet pH measured in 7\/2015 and 7\/2016 NANCY ROSE We are also evaluating our compost and mulch operations and are investigating the potential value of cover crops in our parklike setting. This fall, for example, we are testing the feasibility of growing tillage radish (Raphanus sativus) in the landscape as a no-till method for reducing the soil compaction that has resulted from increased foot traffic and equipment. Tillage radish is an agricultural cover crop known for its long taproot that \"drills\" through the soil, reducing compaction in a natural way; this taproot can grow as long as 6 feet (1.8 meters), and the first 12 to 20 inches (30.5 to 50.8 centimeters) can have a diameter up to 2 inches (5.1 centimeters) (United States Department of Agriculture 2012). When planted in fall, the radishes are winterkilled and decompose, thereby producing large cavities in the soil and releasing nutrients for surrounding plants to take up (United States Department of Agriculture, 2012). If you have been to the Arboretum since midAugust, you may have noticed four 20by 20-foot (6.1-meter) fenced plots in our landscape. In each of these plots we are testing four sowing methods (broadcast seeding, broadcast seeding after aerating the soil, slice seeding, and slice seeding after aerating the soil), with and without a leaf compost cover, to see if we are able to grow this cover crop in an urban, partially shaded setting and, if so, which of these eight methods works best. If successful, we may integrate tillage radish into our management. We intend to continue exploring options and testing them in the landscape, learning from our trials to develop best management practices. Managing the health of our soils to provide the best growing conditions for the over 2,000 diverse taxa that we cultivate from temperate biomes across the world is an essential component to the Arboretum's Plant Health Care program. We hope that our adaptive management approaches will help us develop wellinformed ways to steward the next nearly 400 taxa to join our Living Collections and improve the habitats of our current collections, as well. This tillage radish testing plot is in the Linden (Tilia) Collection. KYLE PORT 12 Arnoldia 74\/2 ? October 2016 (continued from page 8) previous fall comparing siblings were revisited and adjustments were made in ranking based on their overall condition. Once all is checked, lists referred to as \"planting bulletins,\" which include accession numbers, names, and current nursery and final planting locations, are systemically issued to Living Collections Managers once final checks are complete and species' transplanting priority is established. The issuing of a bulletin first triggers the Manager of Plant Records to initiate important database updates and in turn create permanent labels for each plant being transplanted. Before plants leave the production facility, permanent labels are attached and double checked against temporary nursery labels to avoid mix-ups. When those tasks are done, horticulturists are given the green light to start the digging process. Ideally the goal is to complete transplanting before the plants break bud. Taxa such as birch (Betula spp.) and apple (Malus spp.) that leaf out early are the first priority of the digging season and thus will be listed on early bulletins. Others such as ash (Fraxinus spp.) and oaks (Quercus spp.) tend to break dormancy later and can remain in the nursery longer. Containerized plants are the last to be planted as root loss tends to be less severe. The transplanting method known as \"balledand-burlapped,\" or B&B, starts with digging soil out from around the trunk of the plant. As a general rule, the ball radius should be 1 foot (30.5 centimeters) per 1 inch (2.5 centimeters) of trunk diameter. For example, a tree with a 1.5-inch (3.8 centimeter) trunk diameter would have a ball that is 3 feet (91.4 centimeters) across. When digging, larger roots are cut with pruners to avoid tearing, and imperfections in the root system are noted and addressed as needed. Once the ball has been defined and the majority of soil excavated, burlap sheets are placed over the ball and twine is used to hold the ball tightly together to prevent the ball from falling apart and drying out during transplant. When complete, B&B plants are lifted out of the holes and taken to their final planting locations as soon as possible. Arboretum Horticulturist Scott Grimshaw checks planting information for a balled-and-burlapped specimen ready to be moved into the collections. Once on site, the planting hole is dug paying close attention to the height of the ball to avoid making the hole too deep. The root flare, the transition zone between trunk and root system, should be at or slightly above the existing grade and never be covered with soil or mulch since it is a key zone of gas exchange for the plant. Covering the root flare can also lead to the development of a secondary root system and the occurrence of girdling roots. With burlap and twine intact, the ball is placed in the planting hole and final adjustments to planting depth are made, and the tree is viewed from all angles to ensure that the plant is straight. The majority, if not all, of the burlap and twine is then cut away from the ball and the planting hole is backfilled with the excavated soil. A 3- to 4-inch (7.6- to 10.2-centimeter) layer of mulch is applied, making sure not to cover the root flare or trunk, and plants are watered thoroughly to hydrate roots and ensure good soil-to-root contact from ANDREW GAPINSKI KYLE PORT Living Collections 13 Arboretum Horticulturists Scott Grimshaw and Rachel Brinkman lace twine around the burlap covered root ball of a Magnolia amoena (accession 385-2012-A) in preparation for moving it out of the west nursery to its new home along Chinese Path in the Explorers Garden. Arboretum Horticulturists Mark Walkama and Wes Kalloch plant a specimen of black cherry (Prunus serotina, 602-2008B), carefully removing twine and burlap from the root ball before replacing excavated soil. Caring for the Curated Landscape Although much planning and many resources have gone into all phases of collections development from the ANDREW GAPINSKI the start. New plantings are provided with regular watering during their first year of establishment and also in subsequent years when drought conditions occur. Once the transplanting of all accessions on a particular bulletin is complete, the Manager of Plant Records is notified and each plant is visited to collect accurate GPS coordinates. In addition, all temporary marking materials (nursery labels, flagging tape) are removed and permanent labels are repositioned as needed. Watering is key to successful establishment of newly installed accessions. The Arboretum deploys both hand-watering and automated irrigation systems such as this one being installed in the renovated planting area in front of the Hunnewell Building by Horticulturist Greg LaPlume. point of acquisition to establishment on the grounds, the work to preserve and steward these holdings both curatorially and horticulturally has just begun. The Arboretum's curatorial team maps, labels, and regularly inventories and evaluates all accessions, noting such observations as growth, health, damage, and various other metrics. Horticulturally, we seek to keep specimens vigorous and thriving through regular aesthetic and corrective pruning, reduction of weed competition, soil health management, and the evaluation, prioritization, and mitigation of various plant stressors, from pest and disease pressure to drought. With the goal being to maintain the germplasm represented by our collections into perpe- Goats from a local \"goatscaping\" company have been deployed to clear weedy underbrush and invasives such as buckthorn and Oriental bittersweet from tuity, plant production staff sections of the Arboretum. continue to play a key role Dungait, J. A. J., D. W. Hopkins, A. S. Gregory, and A. P. in preserving important lineages through the Whitmore. 2012. Soil organic matter turnover collection of vegetative propagation materials, is governed by accessibility not recalcitrance. such as cuttings and scions for grafting, from Global Change Biology 18: 1781?1796. existing accessions. A lineage may be repropaFriedman et al. 2016. Developing an Exemplary gated because of the decline of a specimen or to Collection: A Vision for the Next Century at create clones for distribution to other instituthe Arnold Arboretum of Harvard University. tions around the world. As we complete our Arnoldia 73(3): 2?18. second year of expeditions for the Campaign for Koller, G. K. 1989. Landscape Curation: Maintaining the the Living Collections, with new lineages and Living Collections Arnoldia 49(1): 65?72 taxa growing in the greenhouses, we anxiously United States Department of Agriculture. 2012. await transplanting the first of the Campaign Oilseed Radish: Raphanus sativus L. Plant material into the Living Collections and the Guide from the USDA Natural Resource Conservation Service. Retrieved 1 August challenges and opportunities that will follow. References Arnold Arboretum. Living Collections Policy. http:\/\/ w w w. a r b o r e t u m . h a r v a r d . e d u \/ p l a n t s \/ collections-management\/living-collectionspolicy\/. Accessed September 20, 2016. Dosmann, M. S. 2015. The History of Minimum Temperatures at the Arnold Arboretum: Variation in Time and Space. Arnoldia 72(4): 2?11. 2016 from: http:\/\/www.nrcs.usda.gov\/Internet\/ F S E _ P L A N T M AT E R I A L S \/ p u b l i c a t i o n s \/ arpmcpg11828.pdf United States Department of Agriculture. 2016. Soil Health. Retrieved 18 August 2016 from: http:\/\/ www.nrcs.usda.gov\/wps\/portal\/nrcs\/main\/ soils\/health\/ Andrew Gapinski is Manager of Horticulture at the Arnold Arboretum. ANDREW GAPINSKI 14 Arnoldia 74\/2 ? October 2016 "},{"has_event_date":0,"type":"arnoldia","title":"Towards Broader Adaptability of North American Deciduous Azaleas","article_sequence":2,"start_page":15,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25613","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d14ea36f.jpg","volume":74,"issue_number":2,"year":2016,"series":null,"season":null,"authors":"Hokanson, Stan C.; Bradeen, James M.; Susko, Alexander Q.","article_content":"Towards Broader Adaptability of North American Deciduous Azaleas Alexander Q. Susko, James M. Bradeen, and Stan C. Hokanson N and many shades in between, providing tremendous spring and summer interest in the garden. Quickly recognized for their horticultural merit, North American deciduous azaleas piqued the interest of plant collectors upon their discovery and continue to be widely lauded by gardeners today. This has led to a proliferation of cultivars and interspecific hybrids that provide a beautiful floral display every year both in gardens and in the wild (Azalea Society of America 2016). WILLIAM (NED) FRIEDMAN orth American deciduous azaleas have long been adored by horticulturists. These plants, which belong to the genus Rhododendron sect. [section] Pentanthera, comprise 15 species distributed from Texas to Florida, extending northwards to southern Maine, and with one species occurring in mountainous areas of Oregon and California. These species display a wide range of flower color, from pure white to deep orange, pink, As a common name, \"rhododendron\" usually refers to large evergreen shrubs whose flowers have ten stamens, such as this elepidote cultivar `Album Grandiflorum' (accession 22810-A) growing in the Arnold Arboretum's Rhododendron Dell. 16 Arnoldia 74\/2 ? October 2016 The Linnaean system classifies organisms into increasingly narrow groups until reaching the individual species level. Classification (to the section level) for the North American deciduous azaleas is shown here (US-GRIN 2016). Kingdom......... Plantae Order............... Ericales Family............. Ericaceae Genus............. Rhododendron Subgenus.........Hymenanthes Section............Pentanthera DANNY SCHISSLER Over 240 unique accessions of Rhododendron sect. Pentanthera exist at the Arnold Arboretum including many interspecific hybrids, various cultivars, and 12 of the 15 deciduous azalea species native to North America (Table 1). The accessions at the Arnold Arboretum have origins in a wide range of environments and could contain many useful adaptations to common abiotic (non-living) stressors. This collection thus represents a valuable asset to woody plant researchers who seek to understand the adaptive potential of these plants to better benefit gardeners and ecosystems alike. Those who have spent any time researching azalea species or cultivars quickly discover a confusing naming system. Azaleas are actually a now informal designation within the genus Rhododendron, a large genus of 1,024 species Azaleas typically have five stamens that are often strongly exserted, as seen on this Alabama azalea (Rhododendron alabamense, 137-2005-E). in the heath family (Ericaceae) with a distribution spanning across the northern hemisphere (American Rhododendron Society 2015). Horticulturists and plant enthusiasts commonly split the genus Rhododendron into two informal groups, \"rhododendrons\" and \"azaleas,\" based on appearance. Rhododendrons are typically considered to be large evergreen shrubs with large, leathery, elepidote (without scales) leaves, or smaller evergreen shrubs with lepidote (with scales) leaves. Rhododendrons are also differentiated from azaleas in flower morphology by the presence of 10 or more stamens (Azalea Society of America 2016). Azaleas, in contrast, are usually considered to be smaller, sometimes deciduous shrubs with pubescent leaves and five to six stamens, though these morphological characteristics can be quite varied within species (Azalea Society of America 2016). The word azalea is derived from the Greek azaleos meaning \"growing in dry soil.\" This name reflected the cliffside and mountain habitats of Rhododendron luteum (yellow azalea), a European species (Lee et al. 1953). However, this is a misleading name because azaleas are generally intolerant of drought and are most commonly found in moist or Table 1: North American deciduous azaleas at the Arnold Arboretum Species Distribution Representative Accession Flowering Details Rhododendron alabamense AL, FL, GA, MS, TN 137-2005 Blooms April-May. Mostly white flowers, some pink coloration. R. arborescens AL, GA, KY, MD, MS, NC, NY, PA, SC, TN, VA, WV 464-81 Blooms June-August. White, fragrant R. atlanticum DE, GA, MD, NC, NJ, PA, SC, VA 108-2007 Blooms April-May; June in northern gardens. White\/ pink, very fragrant. R. austrinum AL, FL, GA, MS 1403-85 Blooms March-April in wild, later in northern gardens. Yellow\/orange, fragrant. R. calendulaceum AL, CT, GA, KY, MD, MS, NC, NY, OH, PA, SC, TN, VA, WV 109-2007 Blooms May-June. Variable color, from pink\/orange\/redorange depending on source. R. canescens AL, AR, DE, FL, GA, KY, LA, MD, MS, NC, SC, TN, TX 468-81 Blooms March-April; May in northern gardens. Pink\/white. R. occidentale CA, OR 21743 (hybrid) R. periclymenoides AL, CT, GA, IL, KY, MA, MD, NC, NH, NY, OH, PA, RI, SC, VA, VT, WV 111-2007 Blooms April-May. Varying shades of pink. R. prinophyllum AL, AR, CT, GA, IL, KY, MA, MD, ME, MO, NC, NH, NY, OH, OK, PA, RI, TX, VA, VT, WV 805-86 Blooms May. Pink. R. prunifolium AL, GA 815-90 Late blooming, JulySeptember. Vivid orange\/red. R. vaseyi NC 1438-85 Blooms April-May. Varying shades of pink. Vulnerable in wild. R. viscosum AL, AR, CT, DE, GA, LA, MA, MD, ME, MO, MS, NC, NH, NJ, NY, OK, PA, RI, SC, TX, VA, VT, WV 112-2007 May-August. White\/pink, depending on germplasm source. Fragrant. AZALEA SOCIETY OF AMERICA 2016 AL=Alabama, AR=Arkansas, CA=California, CT=Connecticut, DE=Delaware, FL=Florida, GA=Georgia, IL=Illinois, KY=Kentucky, LA=Louisiana, MA=Massachusetts, MD=Maryland, ME=Maine, MO=Missouri, MS=Mississippi, NC=North Carolina, NH=New Hampshire, NJ=New Jersey, NY=New York, OH=Ohio, OK=Oklahoma, OR=Oregon, PA=Pennsylvania, RI=Rhode Island, SC=South Carolina, TN=Tennessee, TX=Texas, VA=Virginia, VT=Vermont, WV=West Virginia 18 Arnoldia 74\/2 ? October 2016 the assimilation of the North American deciduous azaleas as a section within the broader subgenus Hymenanthes, which includes many large-leafed evergreen Rhododendron species. Nonetheless, \"azalea\" and \"rhododendron\" remain important horticultural terms for marketing and identification purposes. Rhododendron in the Garden ALEXANDER Q. SUSKO The earliest recorded cultivation of plants in the genus Rhododendron began in the fifteenth century in Japan and involved a cultivar group now known as the Satsuki azaleas, derived from the former Tsutsusi, or subgenus of evergreen azaleas (Callaham 2006). At the time, only the rich and privileged classes in Japan were allowed to grow and possess azaleas as they were deemed \"too beautiful\" for commoners (Callaham 2006). Early botanical explorers such as Robert Fortune in Asia and John Tradescant in colonial North America first brought Rhododendron to prominence through their plant collecting activities in the eighteenth and nineteenth centuries (Cox et al. 1997). They discovered diverse flower color, leaf morphology, and plant habit across the Rhododendron species they encountered. Soon after these plants appeared in European gardens, plantsmen began creating interspecific hybrids. The great diversity in flower colors, fragrance, plant size, ALEXANDER Q. SUSKO mesic forest sites (Hume 1948). Considered a distinct genus (Azalea) until the mid-twentieth century, azaleas were moved into the genus Rhododendron after Sleumer's taxonomy was published in 1949 (Chamberlain et al. 1996). Such taxonomies--the relationships of species based on morphological traits and, increasingly, genetic variation--are complex and frequently revised in a large genus like Rhododendron as new species are discovered. Traditional morphology-based taxonomy uses leaf, bud, floral, and other physical traits to determine relationships between species. However, this kind of physical classification can confuse convergent evolution (evolution of similar adaptations independently across varying taxa) with species similarity, leading to incorrect conclusions about species relationships. Modern molecular-based taxonomy uses differences in DNA sequence to group species based on the likelihood of previously shared ancestry. Since this is based on the timespan back to the most recent common ancestor, the classifications will not be confused by convergent evolution of morphological traits. In the genus Rhododendron, subsequent molecular phylogenies that group species based on genetic variation have shown no distinction between species commonly known as azaleas and rhododendrons (Goetsch et al. 2005). This has led to Swamp azalea (Rhododendron viscosum) has a very wide native range and variable growth habit. Flowers are typically white, as seen on this tagged plant (left) growing near the Fourche la Fave River in the Ouachita National Forest near Y City, Arkansas, but pink flowers also occur, as on this plant (right) found in another population within the Ouachita National Forest, this one near Eagleton, Arkansas North American Deciduous Azaleas 19 KYLE PORT The Ghent Hybrid Azaleas The brightly colored flowers of Ghent hybrid azalea cultivar `Reine des Rouges' (accession 623-61-A) show a resemblance to those of flame azalea (R. calendulaceum), one of the parent species originally used in hybridizing the Ghent azaleas. leaf shape, and leaf pubescence (indumentum) resulting from these efforts led to expanded horticultural interest in rhododendrons and azaleas in the early twentieth century (Hume 1948). Throughout the twentieth century, hobbyists played a major role in the proliferation of new cultivars (Galle 1974). Rhododendron cultivation today spreads across temperate regions of the world, and there are over 28,000 named cultivars in existence (American Rhododendron Society 2016). Deciduous azaleas in sect. Pentanthera are represented by one species each in Europe, Asia, and western North America and 14 accepted species in eastern North America (Towe 2004; Zhou et al. 2008). Deciduous azaleas constitute the largest group of Rhododendron species in North America. Because it holds such a large number of species (compared to other parts of the world), the southeastern United States is considered a major center of diversity for deciduous azaleas (Hume 1953). The great amount of phenotypic variation for flower color, fragrance, The Ghent hybrid azaleas resulted from crosses between the European native azalea R. luteum (yellow azalea) and a number of North American species including R. calendulaceum (flame azalea), R. periclymenoides (pinxterbloom azalea), and R. viscosum (swamp azalea) (Dirr 1998). Initial crosses were made in Ghent, Belgium, starting around 1820, additional hybridization occurred in England, and many Ghent hybrid cultivars were introduced, primarily from Belgium, in the following decades. Although the parentage is quite mixed, these hybrids are often grouped together under the name R. ? gandavense. They are notable for their often fragrant flowers that come in a wide range of colors. The Ghent hybrids lost popularity as other hybrid azaleas were introduced, but 22 Ghent cultivars can still be seen in the Arnold Arboretum collections. and cold hardiness among deciduous azaleas has long generated interest for using North American species in breeding. John Bannister, an English botanist, recorded the first deciduous azalea in North America, Rhododendron viscosum (swamp azalea), in Virginia in 1690 (Galle 1974). Seeds from the newly discovered deciduous azalea species were sent back to Europe where plants were grown for observation beginning in 1734 by the American botanist John Bartram. It was not until 1825 that the first recorded and popular interspecific hybrids were produced using North American deciduous azaleas (Hume 1953) (see textbox above). Although they grow well within their native ranges, lack of cold hardiness and\/or intolerance of high soil pH are major limiting factors to cultivation of these azaleas across much of North America. Relatively few Rhododendron species are native to continental climates, typified by extreme temperature variations throughout the year and characteristic of many regions within the United States. This is reflected in the culti- UNIVERSITY OF MINNESOTA 20 Arnoldia 74\/2 ? October 2016 Research staff member Margaret Gearhart stands next to the original `Northern Lights', a cold-hardy deciduous azalea cultivar, in June 1979 at the Minnesota Landscape Arboretum. vation history of Rhododendron species, which, until fairly recently, were only commonly grown and propagated in the mild and humid climates of Japan, Europe, and the eastern and western coasts of North America. For woody plant breeders in the midwestern United States, developing cultivars for their tolerance to continental climates is of paramount importance for the horticultural success of deciduous azaleas. Fortunately, despite originating from milder climates, individual genotypes from many North American native azalea species have been identified that are exceptionally hardy under extreme temperature variation (Widrlechner 1982). North American deciduous azalea germplasm has undergone relatively little systematic evaluation and had received little breeding focus until the twentieth century, when diversified landscape needs led nursery growers to seek hardier Rhododendron germ- plasm (Hokanson 2010). For example, suburbanization in the United States following World War II led to an increased need for landscaping plants to beautify new developments (Whitehand and Larkham 1992). During the middle of the twentieth century, few ornamental plants adapted to the upper midwestern United States existed (UMN Fruit Breeding Farm 1954). Later, University Agricultural Experiment Stations in states like Minnesota began funding breeding programs for ornamental plants to develop and introduce promising cultivars for landscape use (Widrlechner 1982). Many ornamental shrubs were trialed at the time on the grounds of the newly founded Minnesota Landscape Arboretum, including several deciduous azalea species and interspecific hybrids developed previously in Europe. After years of parental and seedling testing, plants from crosses made in 1957 between mollis azaleas (R. ? kosteranum), North American Deciduous Azaleas 21 which are a group of Asian hybrid deciduous azaleas, and the American species R. prinophyllum proved to be hardy to at least -35?C (-31?F) (Johnson and Snyder 1966). One seedling from this breeding population later became the cultivar `Northern Lights' and was released in 1978 as part of the Northern Lights Series, the first flowering azaleas bred for midwestern climates (Pellett and Vos 1978). The University of Minnesota woody landscape plant breeding program has maintained the longest standing breeding program for deciduous azaleas in North America, and continues to actively release cultivars under the Northern Lights Series today (Widrlechner 1982; Hokanson 2010; Hokanson et al. 2015). Identifying Adaptive Genetic Variation NANCY ROSE Though certain stress tolerances (cold hardiness in particular) of azaleas within the University of Minnesota breeding program are well documented, understanding of trait variation for other abiotic stresses in natural Rhododendron sect. Pentanthera populations remains limited. Tolerance to abiotic stressors such as winter cold, summer heat, soil pH, and soil salinity are necessary for successful plant growth both in natural and cultivated settings. Abiotic stress tolerance in woody species like deciduous azaleas can arise genetically through humanmediated or natural selection. However, a lack of understanding about the nature of adaptive genetic variation for such traits is not unique to azaleas; a knowledge gap is common across woody plants and impedes understanding of their maximum adaptation in both horticultural and ecological contexts. Genetic variation, or variation in the DNA sequences between individuals in a species, is often a major factor in influencing the adaptation of an individual to an environment. In situations when individuals possessing certain patterns of genetic variation reproduce more successfully under an environmental stress, genetic variation is An azalea breeding trial bed at the University of Minnesota Horticultural Research Center includes introduced cultivars `Lemon Lights' (left) and `Northern Hi-Lights' (foreground) as well as numbered selections. ALEXANDER Q. SUSKO 22 Arnoldia 74\/2 ? October 2016 Open longleaf pine (Pinus palustris) forest is a typical habitat for Rhododendron viscosum subpopulations in the Apalachicola National Forest near Tallahassee, Florida. considered to be adaptive. Adaptive genetic variation and associated traits are inherited by offspring; identifying such traits is an important (but time consuming) part of breeding all plants. The development of the Northern Lights Series is a prime example: it took 21 years from evaluating initial parents, making crosses, and field testing the seedlings before a reliably cold hardy cultivar was introduced. This is largely due to the biological constraints of breeding woody perennial species, including long juvenile periods (time prior to sexual maturity of the plant) that slow the breeding process. This also complicates efforts to understand the genetics behind important traits, further hindering the discovery of potentially adaptive genetic varia- tion. It is imperative to understand such genetic variation in breeding populations if horticultural breeders wish to sustain the development of new, adapted cultivars. New research technologies present opportunities to more efficiently identify adaptive genetic variation for abiotic stress tolerance in populations of North American deciduous azaleas with important implications for cultivar development. Linking genomic sequence and environmental data, an emerging approach known as landscape genomics, makes it possible to identify genetic variation in plant populations that depend on an environmental factor (Rellstab et al. 2015). It is through this framework that genetic variation is declared North American Deciduous Azaleas 23 ALEXANDER Q. SUSKO Arboretum is currently being used to test these landscape genomic approaches by obtaining environmental data from the origin points of the accessions and performing genetic sequencing on those accessions. The goal is to identify potentially adaptive DNA variation in North American deciduous azaleas that could benefit future breeding efforts by identifying promising species or populations as parents for future cultivar development. For example, patterns of genetic variation might be detected when looking at a set of individual azaleas originating from environments with varying degrees of average winter minimum temperatures. If an association is detected between patterns in the genetic variation and average winter minimum temperature, it is possible that the genetic variation confers an adaptive advantage in those ALEXANDER Q. SUSKO as potentially adaptive. Although some follow-up experiments are necessary to confirm the adaptive nature of this genetic variation discovered using this approach, this advancement gives scientists and breeders a new way to quickly screen diverse wild germplasm to identify populations that display unique adaptations to the environment. Collections at the Arnold Arboretum and our program's prior sampling of wild populations of swamp azalea constitute an ideal source of plants to help answer these types of research questions. Most importantly, plants in this collection are linked to detailed information about their collection locations that allow for relationships between genetic and environmental variation to be investigated. North American deciduous azalea germplasm at the Arnold Rhododendron austrinum (orange azalea; accession 1403-85) is native primarily to the Florida panhandle and adjacent regions in Mississippi, Alabama, and Georgia. It bears fragrant yellow and orange flowers. Rhododendron vaseyi (pinkshell azalea; accession 657-70) is rare in the wild, with only a few known populations, all in North Carolina. The Arnold Arboretum has a number of conservation accessions of this beautiful pink-flowered azalea. KYLE PORT 24 Arnoldia 74\/2 ? October 2016 WILLIAM (NED) FRIEDMAN Rhododendron prunifolium (plumleaf azalea) is a late-season (July to September) bloomer, bearing flowers in shades of red and orange. The flowers of Arnold Arboretum accession 815-90-J, seen here, are an especially vibrant cherry red. Rhododendron calendulaceum (flame azalea; accession 109-2007-J) is notable for its brightly colored flowers that range from yellow to orange to bright red. This species is native to the Appalachian Mountains ranging from New York to Georgia. NANCY ROSE North American Deciduous Azaleas 25 The flowers of Rhododendron periclymenoides (pinxterbloom azalea; accession 3237-C) range from white to pink to purplish pink. It grows in mesic forests and wetlands from New Hampshire and Vermont to Alabama. KYLE PORT 26 Arnoldia 74\/2 ? October 2016 NANCY ROSE Rhododendron atlanticum (coastal or dwarf azalea; accession 108-2007-A) is native to coastal regions in the Mid-Atlantic and Southeast. Its white to light pink flowers are sweetly fragrant. Rhododendron prinophyllum (roseshell azalea) has a wide native range from Maine to Georgia and Texas. Its flowers may be light to deep pink. North American Deciduous Azaleas 27 azalea populations. While existing deciduous azalea germplasm has been well characterized for cold hardiness within the University of Minnesota breeding program, many other sources of tolerance to abiotic stressors (e.g., heat, temperature variability, drought) remain to be identified within this group of plants. Such variation is of great interest to breeders seeking to improve stress tolerance through breeding. The approaches introduced above could be extended in the future towards informing conservation efforts that leverage the adaptive potential of populations. For the existing deciduous germplasm at the Arnold Arboretum, this research will characterize the collection for its adaptive potential to common environmental stressors. Such efforts could greatly inform future collection and conservation efforts for these species should any generate breeding or conservation interest in the future. Ultimately, we hope that this approach will further enable the cultivation of these North American horticultural gems in as many landscapes as possible. Literature Cited A m e r i c a n R h o d o d e n d r o n S o c i e t y. h t t p : \/ \/ w w w. rhododendron.org\/. Last accessed January 2016. Azalea Society of America. http:\/\/azaleas.org\/. Last accessed January 2016. Callaham, R. Z. 2006. Satsuki Azaleas for Bonsai and Azalea Enthusiasts. Passumpsic, Vermont: Stone Lantern Publishing. Chamberlain, D., R. Hyam, G. Argent, G. Fairweather, and K. S. Walter. 1996. The genus Rhododendron: its classification and synonymy. Edinburgh, Scotland: Royal Botanic Garden Edinburgh. Cox, P. A. and K. N. Cox. 1997. The Encyclopedia of Rhododendron Species. Perth, Scotland: Glendoick Publishing. Dirr, M. A. 1998. Manual of Woody Landscape Plants (5th ed.). Champaign, Illinois: Stipes Publishing LLC. Galle, F. C. 1974. Azaleas. Birmingham, Alabama: Oxmoor House, Inc.. Goetsch, L., A. J. Eckert, B. D. Hall, and S. B. Hoot. 2005. The molecular systematics of Rhododendron (Ericaceae): a phylogeny based upon RPB2 gene sequences. Systematic Botany 30:616?626. Hokanson, S. C. 2010. \"Lights\" in the land of 10,000 lakes. In: Rhododendrons, camellias and magnolias. London: Royal Horticultural Society. Hokanson, S. C., S. T. McNamara, N. Rose, K. Zuzek, and H. Pellett. 2015. `UMNAZ 493' and `UMNAZ 502'; Two new cold hardy deciduous azalea selections from the University of Minnesota. HortScience 50(9)S:053 (Abstr.). Hume, H. H. 1948. Azaleas: Kinds and Culture. New York: The MacMillan Company. Hume, H. H. 1953. Azaleas and Camellias (Revised). New York: The MacMillan Company. Johnson, A. G. and L. C. Snyder. 1966. Breeding azaleas for Minnesota. Quarterly Bulletin of the American Rhododendron Society 20:163?165. Lee, F. P., F. O. Coe, B. Y. Morrison, M. Perkins, and F. Weiss. 1953. The Azalea Handbook. Baltimore: Monumental Publishing Company. Pellet, H. and F. Vos. 1978. Northern Lights. Minnesota Agricultural Experiment Station, Miscellaneous Report #155. Rellstab, C., F. Gugerli, A. J. Eckert, A. M. Hancock, and R. Holderegger. 2015. A practical guide to environmental association analysis in landscape genomics. Molecular Ecology 24(17): 4348?4370. Towe, C. L. 2004. American Azaleas. Portland, Oregon: Timber Press. United States National Plant Germplasm System (GRIN), https:\/\/npgsweb.ars-grin.gov\/gringlobal. Last accessed June 2016. University of Minnesota Fruit Breeding Farm. 1954. Report of the Fruit Breeding Farm Visitors Committee. Minnesota Horticulturist, 82(9):143. Whitehand, J. W. R. and P. J. Larkham. 1992. Urban Landscapes: International Perspectives. Hove, East Sussex, United Kingdom: Psychology Press. Widrlechner, M. 1982. Studies on the Breeding Potential and Genetics of Hybrid Azalea, Rhododendron ? Kosterianum Schneider ? Rhododendron Prinophyllum (Small) Millias. St. Paul, Minnesota: University of Minnesota Doctoral Thesis. Zhou, W., T. Gibbons, L. Goetsch, B. Hall, T. Ranney, and R. Miller. 2008. Rhododendron colemanii: A New Species of Deciduous Azalea (Rhododendron section Pentanthera; Ericaceae) from the Coastal Plain of Alabama and Georgia. Journal American Rhododendron Society 62(2): 72?78. Alexander Susko is currently a PhD student in the Applied Plant Sciences graduate program studying plant breeding and molecular genetics, James Bradeen is a professor and department head in the Department of Plant Pathology, and Stan Hokanson is a professor of woody plant breeding and genetics in the Department of Horticultural Science, all at the University of Minnesota, Twin Cities. "},{"has_event_date":0,"type":"arnoldia","title":"BOOK EXCERPT: Urban Forests: A Natural History of Trees and People in the American Cityscape","article_sequence":3,"start_page":28,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25610","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170856d.jpg","volume":74,"issue_number":2,"year":2016,"series":null,"season":null,"authors":"Jonnes, Jill","article_content":"In Urban Forests: A Natural History of Trees and People in the American Cityscape, author Jill Jonnes presents stories from the early days of the urban tree movement on through the challenges that face urban forests today. Excerpted here is Chapter Seven from the book. Book Excerpt 29 30 Arnoldia 74\/2 ? October 2016 Book Excerpt 31 32 Arnoldia 74\/2 ? October 2016 Book Excerpt 33 34 Arnoldia 74\/2 ? October 2016 Book Excerpt 35 Excerpt from Urban Forests copyright Jill Jonnes published September 27, 2016, by Viking, an imprint of Penguin Publishing Group, a division of Penguin Random House LLC. ISBN: 9780670015665 http:\/\/www.penguinrandomhouse. com\/books\/314478\/urban-forestsby-jill-jonnes\/9780670015665\/ "},{"has_event_date":0,"type":"arnoldia","title":"Gray's Bird Cherry (Prunus grayana): A Fitting Tribute to Asa Gray","article_sequence":4,"start_page":36,"end_page":37,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25611","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d1708926.jpg","volume":74,"issue_number":2,"year":2016,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Gray's Bird Cherry (Prunus grayana): A Fitting Tribute to Asa Gray Michael S. Dosmann I f you've ever noticed the similarity between North American and Asian species of Liriodendron, Hamamelis, or Stewartia, then you've recognized the same biogeographic phenomenon that botanists have for well over a century (for an overview of disjunct floras see: Yih, D. 2012. The Eastern Asian?Eastern North American Floristic Disjunction. Arnoldia 69(3): 14?22). One of the first to recognize the similarity between species native to eastern North America (ENA) and eastern Asia (EA), specifically Japan, was Harvard botany professor Asa Gray, who first compared the two floras in 1840 while reviewing Siebold and Zuccarini's Flora Japonica. Fascinatingly, much of Gray's own evolution of thought related to biogeography was nurtured by his extensive correspondence with Charles Darwin. Gray's work recognized \"identical species\" that co-occurred in ENA and EA, and in an 1859 analysis he referred to a Japanese bird cherry as either Prunus virginiana (of North America) or possibly P. padus (a Eurasian species), though his inclination was that it was P. virginiana. Dutch botanist Friedrich Miquel described this bird cherry as P. padus var. japonica in 1865, but it was Russian botanist Carl Maximowicz who, in 1883, named it as a separate species, P. grayana, commemorating Asa Gray. Prunus grayana along with P. virginiana, P. serotina, and P. padus form a group of deciduous, racemose-flowering Prunus species that have a compelling evolutionary and geographic history that extends from eastern Asia to eastern North America and northern Eurasia. How fitting a specific epithet! I was first drawn to Gray's bird cherry in the spring of 2001 while ogling spectacular floral displays in the Arboretum's Living Collections. The 2- to 4-inch (5- to 10-centimeter) -long terminal racemes of densely packed white flowers (each about 1\/3 inch [7 to 8 millimeters] across) have leafy bases and lack peduncles: the lowermost flower emerges directly from a leaf axil. The abundant flowers typically appear in early to mid-May, turning the entire canopy into a dazzling green and white display that lasts about a week or two. The fruits mature from green to bright red and eventually a dark purple by mid-August. The elliptic leaves, up to 4 inches (10 centimeters) long, become intensely yellow gold in the autumn, while the short petioles and midveins often turn bright red--quite a striking contrast. Mature trees are rounded to upright and can grow nearly 50 feet (about 15 meters) tall. Although uncommon in cultivation, P. grayana offers wonderful spring and autumn ornamental interest. Arboretum specimens have been free of major disease or pest issues. It grows at the Morton Arboretum, near Chicago, confirming USDA Zone 5 cold hardiness, and also survives the oppressive heat and humidity at Philadelphia's Morris Arboretum and the United States National Arboretum in Washington, D.C. I have never noticed naturalized seedlings, but since birds eat the fruit and may disperse seeds, I recommend careful invasiveness evaluation before major production and introduction into the trade. Prunus grayana is native to Japan, where it is common in mesic forests from Hokkaido to the southern islands of Shikoku and Kyushu, and also occurs in mainland China from Zhejiang westward to Yunnan. The Arboretum currently has eight living specimens, all from Japan. The oldest (16694-A) was collected by E. H. Wilson in 1914 on Mount Hayachine in northeast Honshu. It grows near Forest Hills Gate, stands 35.4 feet (10.8 meters) in height, and has a diameter of 30.3 inches (77 centimeters) at 2 feet (0.6 meters) above the ground. Our tallest tree, at 42.3 feet (12.9 meters), is 1698-77-C, which grows nearby with three others (1191-77-A and B, and 1698-77-B). All were collected on Hokkaido by then Arboretum staff members Richard Weaver and Steven Spongberg during their 1977 expedition. They also made an additional collection (1777-77) from northern Honshu, with specimens A and C now growing on Bussey Hill and specimen D on Peters Hill. So, this autumn (or during bloom next spring), take a tour of our collection of Gray's bird cherries and think about Asa Gray's laudable contributions to botany-- especially his notice of the North AmericanAsian floral connection. Michael S. Dosmann is Curator of Living Collections at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23452","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd14ea328.jpg","title":"2016-74-2","volume":74,"issue_number":2,"year":2016,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"A Concise Chronicle of Propagation","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25605","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170b76d.jpg","volume":74,"issue_number":1,"year":2016,"series":null,"season":null,"authors":"Alexander III, John H.; Enzenbacher, Tiffany","article_content":"CAMPAIGN FOR THE LIVING COLLECTIONS The Campaign for the Living Collections kicked off last fall with plant collecting trips to China and Idaho, which Curator of Living Collections Michael Dosmann and Manager of Plant Records Kyle Port wrote about in the last issue of Arnoldia. Once newly acquired fruits, seeds, cuttings, divisions, and plants arrive at the Arboretum, the production staff at the Dana Greenhouses takes over. In this issue, Manager of Plant Production Tiffany Enzenbacher and Plant Propagator John H. Alexander III describe the process of shepherding new accessions from the greenhouse bench to final production nurseries, the last step before plants move to a permanent location on the Arboretum grounds. A Concise Chronicle of Propagation Tiffany Enzenbacher and John H. Alexander III P lant propagation historically has been recognized as an integral component of the Arnold Arboretum's mission. In fact, the Arboretum's second employee (inaugural director Charles S. Sargent was the first) was plant propagator Jackson Dawson, hired in 1873, the year after the Arboretum was established. Since Dawson was Sargent's only employee, he served not only as the propagator but also the superintendent (Geary and Hutchinson 1980) and remained with the Arboretum until his death in 1916. The Arboretum's other long-term and influential propagators--William H. Judd (employed from 1913 to 1946), Alfred J. Fordham (1929 to 1977), and John H. Alexander III (1976 to 2016)--and shorter-term propagators followed suit after Dawson. Through horticultural expertise, experience, and old-fashioned trial and error, they coaxed seeds to germinate and cuttings to grow roots, successfully propagating taxa novel to New England and North America. The propagation facilities have moved five times over almost a century and a half and have seen many exciting horticultural accomplishments by Arboretum propagators and production staff. In 1873, the Arboretum shared growing space with the Bussey Institution, then relocated in 1886 to a small land plot and 20-foot by 50-foot greenhouse on the property at 1090 Centre Street, where Dawson then resided (Geary and Hutchinson 1980). These modest accommodations were soon outgrown and a new greenhouse was constructed on Orchard Street across from the Arboretum (off of the Arborway) in 1917 (Howard 1962). With intensified Arborway traffic and road widening, production was moved back to land adjacent to the Bussey Institution in 1928. As additional space needs arose, along with the desire for a more up-to-date building, ground broke to construct the Charles Stratton Dana Greenhouses in 1961. The donation for the Greenhouses was provided by Mrs. William R. Mercer (n?e Martha Dana), and was named in honor of her father, Charles S. Dana (Howard 1962). This complex houses the present facilities, including specialized equipment and environments for seed, cutting, and grafting propagation, greenhouse and outdoor bench space for containers, an evaluation ARCHIVES OF THE ARNOLD ARBORETUM The Arnold Arboretum's greenhouse at the Bussey Institution was built in 1928. This view is of the greenhouse interior in 1949, photographed by Heman Howard. Views of the Dana Greenhouses in spring 1966 by Heman Howard (above) and again in fall 1974 by Alfred J. Fordham (below). TIFFANY ENZENBACHER ARCHIVES OF THE ARNOLD ARBORETUM Propagation 3 Recent renovations to the Dana Greenhouses increase efficiency and save water and energy. The seed propagation house is now equipped with an improved mist system that features hanging mist assemblies, which allows for maximum use of bench space. Other revamped features include sectioning the house into three irrigation zones with as many isolation valves in each zone, which allows for the flexibility to tailor water needs to specific taxa. Mist frequency can be controlled to come on at intervals of 2 to 180 minutes, with the duration of a mist event ranging from 2 to 60 seconds. LED (lightemitting diode) lighting was installed in fall 2015. The LumiGrow Pro 325 LEDs utilize 70% less energy than our previous HID (high intensity discharge) lamps and produce 70% less heat. Lights are used to extend the day length during short days. nursery, three longer-term nurseries, a cold storage building for overwintering containers, and the Bonsai and Penjing Pavilion. Through facility relocations and many staff changes in the years since the Arboretum's inception, plant propagation and production have remained center stage. The current tenyear Campaign for the Living Collections (Friedman et al. 2016), which focuses on acquiring nearly 400 wild-collected plant taxa, will assuredly keep propagation in the limelight well into the future. The Campaign's list of desiderata features taxa selected because they increase the phylogenetic and biological breadth of Arboretum collections, belong to geographically disjunct clades, are marginally hardy or threatened in the wild, or can be used to create a \"living type specimen\" in genomic research. Last September, the Dana Greenhouses staff received 100 new accessions (seeds, cuttings, plants) from expeditions related to the Campaign. Seeds from many accessions have already germinated, and others such as paperbark maple (Acer griseum) may take several cycles of warm and cold stratification to germinate uniformly. We look forward to transitioning individuals through the phases of production here at the Dana Greenhouses, with the end goal of having plants in their permanent locations in the living collections for researchers to study, children to learn from, and the public to enjoy. PROPAGATION MATERIAL ARRIVES In autumn, as plants in the living collections are slowing in growth and their foliage begins to abscise, the \"growing season\" in the Dana Greenhouses is just commencing. Production staff is overwhelmed with anticipation about what seeds, fruit, cuttings, and plants we will be receiving from foreign and domestic expeditions. However, once the highly sought-after fruit or cutting has been harvested from its parent and is now at long last in the hands of an Arnold Arboretum explorer, its trip to the Arboretum's greenhouses is nowhere near complete. As Curator of Living Collections Michael Dosmann and Plant Records Manager Kyle Port (2016) explained in the last issue of Arnoldia, the United States Department of Agriculture's Animal and Plant Health Inspection Service KYLE PORT 4 Arnoldia 74\/1 ? August 2016 Fleshy fruits like these from wild-collected Washington hawthorn (Crataegus phaenopyrum) are soaked, rubbed, and sieved to separate the pulp from the seeds. (APHIS) requires a specialized permit to import foreign seeds into the United States. This permit allows for the importation of a small quantity of seeds, pending a successful evaluation for hitchhikers--noxious weed or parasitic plant seeds, insect pests, or pathogens of concern. The Arboretum typically has seeds routed to the APHIS Plant Protection and Quarantine inspection station at John F. Kennedy International Airport in Jamaica, New York. Because the several day to weeklong inspection process is so complex and vital, foreign seeds have to be clean (removal of fruit surrounding seeds), properly labeled, and limited to only 50 seeds (or 10 grams [0.35 ounces]) per package. Should the scrutinizing agent discover any unwanted travelers on the coveted soon-to-be Arboretum seeds, the entire content of the package fails to pass the test, and the voyage for that seed lot ends there. Because much time is spent to meticulously clean the seeds and package them correctly in the foreign country, the majority pass through inspection and are then shipped on to the Arboretum, where the true journey through Clockwise from upper right: Curator of Living Collections Michael Dosmann opens a shipment of seeds from the APHIS Plant Protection and Quarantine inspection station at John F. Kennedy International Airport in Jamaica, New York. The seeds were acquired during the North AmericaChina Plant Exploration Consortium (NACPEC) expedition in September 2015 and were sent directly from China to the inspection station. Jack Alexander prepares to sow Quercus rehderiana acorns in individual containers. These acorns were shipped directly to the Arboretum from collaborators in China. We then sent them to APHIS for inspection, and, after passing examination, they were returned to the Arboretum where they were cold stratified to break dormancy. Kyle Port collected this bunchberry accession (Cornus canadensis 209-2015) as a whole plant during his expedition to northern Idaho in September 2015. ALL PAGE 5 PHOTOS BY TIFFANY ENZENBACHER This paperbark filbert (Corylus fargesii) seedling is identified with accession number, form received as (SD = seed), and collection information on both hand-written and thermalprinted labels. Sixteen seeds were acquired during the 2015 NACPEC expedition and so far three have germinated. Seed Propagation Seeds from our own staff collectors, collaborators, and other gardens never arrive in those colorful packets seen on garden center display racks. Our seeds may arrive in small, resealable polyethlene bags, coin envelopes labelled in beautiful cursive writing, or sheets of paper neatly folded into packets. All will be carefully handled as they enter the propagation process. The first step is examination, since occasionally those packets contain more than seeds. Fruit remnants, cones, and chaff may arrive with the seeds, plus the occasional weevil or other insect. Collections made in foreign countries are thoroughly cleaned before being shipped since they will have to pass an inspection by APHIS (see page 4). Collections made within the United States by our own staff are seldom cleaned before being shipped back to the Arboretum, so at the greenhouse we often get to unpack boxes full of polyethylene bags containing rotting and fermenting fruits. Seeds from other arboreta and botanic gardens, be it foreign or domestic, are usually neatly cleaned and packaged. Not every seed in every packet will germinate, though. We once obtained a half kilogram (about a pound) of wild-collected Chinese sweetgum (Liquidambar acalycina) seeds, but ended up with only a tablespoon of viable seeds while the rest were undeveloped. Anyone unfamiliar with sweetgum seeds could easily make this mistake since sweetgum fruits often hold more undeveloped seeds than sound seeds. Careful visual inspection may help determine sound from unsound seeds, but not always. For example, bald cypress (Taxodium distichum) seeds are not uniform and could easily be tossed out with the cones. Before sowing, plant propagators routinely remove all that is not seed (e.g., fruit pulp, capsules, cones) because it is likely to host fungi, attract insects or rodents, or, in the case of fruit pulp, inhibit seed germination. Cleaning may involve soaking, drying, sieving, or a combination of these and other techniques. Sometimes seeds and chaff are all so tiny, and separating the two so difficult, that it only makes sense to clean reasonably well and sow it all. Freshly collected seeds generally germinate in higher percentages than stored seeds so we go to work quickly once seeds arrive. Before sowing the cleaned seeds we need to know the best protocol for germination for that particular species. For many plants, past experience or a search of seed propagation reference materials provides well-established protocols for germination variables such as soil temperature, day length, or light\/dark requirements. Seeds of most temperate zone species require cold stratification, which simulates winter conditions, and will germinate in higher percentages if they first experience 30 to 120 days at temperatures just above freezing. We routinely place seeds into polyethylene bags containing a moist, well-drained medium and refrigerate at 40?F for 90 days. The seeds of some species need both warm and cold stratification periods. Examples include paperbark maple (Acer griseum) and related trifoliate maples, the dove tree (Davidia involucrata), and most viburnums (Viburnum spp.). And there are also many species whose seeds don't strictly require cold stratification (heath family [Ericaceae] members, for example) but they germinate more uniformly and in higher percentages if first given a one month cold stratification so we often opt for that treatment. Another obstacle for germination in some seeds is the presence of an impermeable seed coat. Plants in Fabaceae, the pea family, often have impermeable seed coats, so we typically scarify seeds of any fabaceous species, whether known or new to us, by rubbing on sandpaper or a file. Scarified seeds are then soaked in water; if they \"imbibe\" and swell to about twice their size, they are ready to be sown or stratified. For all seeds, imbibition is the first step in germination (and why garden seed packets always exhort gardeners to \"keep soil moist after sowing\"). KYLE PORT Keeping records is an essential part of plant propagation. To track germination percentages and successful protocols, we count seeds (or make a close estimate) before they are sown. Once the number of seeds is known and a protocol has been determined, we begin the specified treatment. With species that haven't been grown before at the Arboretum or for which no established protocol can be found, we may experiment and try a variety of treatments if there are plenty of seeds. If there are only a few seeds, we rely on experience and best judgement to pick a treatment. Once stratifications (if needed) are complete, seeds are sown in flats and placed in a warm, humid greenhouse with the option of supplemental lighting. The best time to sow seeds is in the early spring but that timing isn't always possible, so supplemental lighting allows us to lengthen the photoperiod to simulate the longer days of spring and summer. When seedlings reach sufficient size they are potted up in individual containers, ready to continue through our production system. Modern technology has changed many greenhouse peripherals--we now use LED lights, thermostats, soil heating mats, and precise irrigation--but nature's requirements for seed germination haven't changed, and we accomplish that in much the same way as did the Arboretum's earliest propagators. Many accessions of fruits and seeds are processed at the Dana Greenhouses. When it's large enough, this healthy Rosa moyesii seedling will be planted in the Shade House. TIFFANY ENZENBACHER the production system begins. On occasion, this step is skipped and seeds are shipped directly to the Arboretum from a foreign country. Since the inspection process is required by law and is essential in mitigating the introduction of invasive and\/or threating agents to agriculture and the environment, greenhouse staff sends the material to APHIS to be inspected prior to any germination treatment. If domestic fruits (berries, capsules, samaras, etc.), cuttings, or plants are acquired, such as materials that Kyle Port collected on his expedition to Northern Idaho last fall, they are shipped directly to the greenhouse. It should be noted that obtaining material from expeditions is not the sole means by which the greenhouse procures plants. Propagules and plants are also obtained by several other methods: through Index Seminum (seed list) exchanges offered by botanical institutions, from other gardens or arboreta, or by purchasing from nurseries (particularly when acquiring cultivars). However, upon receiving any new seed, cutting, or plant, no matter what it is or where it is from, the first step that production staff takes on is accessioning. Similar to all museums, the Arboretum has a number classification system in place so that each plant can be treated as a specimen with a unique, recognized background. The accession number is composed of a number-year unit. For example, the number 2742015 signifies the 274th plant material lot received in 2015. For every accession, abbreviations such as SD (seed), CT (cutting), PT (plant) denote the form of material received. TIFFANY ENZENBACHER 8 Arnoldia 74\/1 ? August 2016 MOVING UP After a seedling has rooted into its growing container, the next phase through the production system beckons. The Shade House, true to its name, is covered by woven polyethylene fabric that Seedlings of Acer oblongum, a semi-evergreen maple native to the sub-Himalayan region, wait in the outdoor container area before being transplanted into the Shade House in 2016. The first Arboretum accession of this species in 1908 comprised seeds collected by Ernest H. Wilson in China. This most recent accession, 272-2015, represents the sixth accession of A. oblongum grown at the Arboretum. One accession in particular that we are eager about having the opportunity to move through the production system is Moyes rose (Rosa moyesii). First collected by Antwerp E. Pratt in 1893, R. moyesii was introduced from Western Sichuan in 1903 by Ernest H. Wilson, Arboretum plant explorer and botanist, and William Botting Hemsley. Wilson collected R. moyesii on the Tibetan frontier, near Tatien-lu, while on expedition for James Veitch and Sons Nurseries (Wilson 1906). Wilson noted that \"the species is not uncommon in shrubberies on the mountains between Mt. Omi and Tatienlu,\" and described the solitary flowers as \"very dark red ... 5 to 6.5 cm across\" and \"singularly pleasing.\" Wilson wrote that R. moyesii was \"named in compliment to the Rev. James Moyes, of the China Inland Mission, stationed at Tatien-lu, to whom I am much indebted for hospitality, assistance, and companionship on one long An illustration of Rosa moyesii from the October 21, 1916, issue of The Garand interesting journey in den, a weekly gardening journal published in London from 1871 to 1927. Eastern Tibet.\" Sargent later commissioned Wilson to collect for the Arboretum, and in 1909 Wilson was successful in acquiring seeds--the Arboretum's second accession of R. moyesii (17091). The first accession (6827) was obtained two years prior, as a plant, directly from Veitch Nurseries. The blossoms of R. moyesii are unique, an intense deep red. Wilson wrote in 1930, \"few if any wild species of Rose have created so much interest as this native of the ChinoThibetan borderland.\" However, he also noted that \"unfortunately, in this climate the flowers bleach rapidly and New England gardens will never know the real beauty of this Rose,\" which prompted him to add that the \"hips ... in this country are more attractive than its flowers.\" The showy orangish red hips have an elongated, bottle-like shape and can reach 2 inches (5 centimeters) long. R. moyesii is still a popular species rose today, but `Geranium', a selection introduced to North America by the Arboretum, is more widely grown. `Geranium' was written about in 1960 by Donald Wyman, Arboretum horticulturist from 1935 to 1970, as a plant of possible merit. It is more compact than the species, with larger hips. This selection originated at the Royal Horticultural Society's garden at Wisley in southern England. ARCHIVES OF THE ARNOLD ARBORETUM A Rose Returns to the Arboretum JENNIFER RIOUX 10 Arnoldia 74\/1 ? August 2016 allows only 45% of the light to pass through. This keeps the vulnerable plants less stressed after transplant. Seedlings and small cuttings or plants are transplanted into the highly organic soil of this evaluation nursery in late spring to early summer and are well tended throughout the season. Plants are mulched in and hand watered until established. There is an overhead sprinkler system for irrigating the entire nursery when necessary. Rodents have been problematic, occasionally damaging all individuals within an accession, so caging plants that they appear to be most attracted to such as horse chestnut (Aesculus), hickory (Carya), and oak (Quercus) has become mandatory in recent years. The Shade House also offers a first test of cold hardiness. Since the vast majority of Arboretum Former Isabella Welles Hunnewell 2014 Intern and Term Employee Olivia Fragale (left) and former Hunnewell 2015 Intern Carly Troncale (right) standing next to cages in the Shade House that they constructed to protect seedlings from possible rodent damage. TIFFANY ENZENBACHER Manager of Plant Production Tiffany Enzenbacher rearranges flats in the center alley to make room for Cornus sericea (accession 257-2015) and Rosa moyesii (accession 285-2015) seedlings. All seedlings transition from the greenhouse to the outdoor container area before being transplanted into the Shade House. Propagation 11 HEADING TO THE COLLECTIONS After the individuals in an accession are large enough to transplant, shrubs get containerized and trees continue their journeys through the facility into one of three longer term nurseries. After entering the production facility as a propagule, trees take anywhere from five to seven years in the system before they are robust enough to be transplanted into the living collections. Shrubs are at the greenhouse for three to five years on average. The voyage of an accession through the Dana Greenhouses concludes when the individuals are planted into their sited location out on the grounds. Now a new, much longer passage of life begins. KYLE PORT accessions funnel through here, and because the greenhouse area is in a recognized cooler microclimate of the Arboretum (Dosmann 2015), it provides a rudimentary assessment of hardiness. However, if a species is known to be marginally hardy, one to several individuals may be containerized instead of being planted in the Shade House. Those individuals would then subsequently be planted in a warmer microclimate of the Arboretum to increase their likelihood of survival during typical Zone 6 (average annual minimum temperatures -10 to 0?F [-23.3 to -17.8?C]) Boston winters. Along with hardiness, seedlings are also evaluated for form and vigor. Shrubs and trees gain size in the container area (foreground) and the East Nursery (beyond container area). Curator Michael Dosmann, Manager of Horticulture Andrew Gapinski, and Manager of Plant Production Tiffany Enzenbacher regularly walk through the nurseries and container areas and determine which individuals will be designated for upcoming plantings. TIFFANY ENZENBACHER Young trees may grow for several years in the well-mulched East Nursery adjacent to the Dana Greenhouses. Once they attain sufficient size, they will be transplanted to permanent locations on the Arboretum grounds. Long-time Arboretum plant propagator Jack Alexander was sowing seeds in the Dana Greenhouses earlier this year. Divisions of twinflower (Linnaea borealis, 198-2015) collected by Kyle Port during last year's North Idaho Expedition take root under mist. This trailing, semi-woody evergreen has a wide circumboreal distribution and was named in honor of famed Swedish botanist Carl Linnaeus. KYLE PORT TIFFANY ENZENBACHER Clockwise from upper left: TIFFANY ENZENBACHER Propagation 13 The cold storage building provides a controlled climate during winter for dormant seedlings, rooted cuttings, containerized plants, and the bonsai and penjing collection. The Campaign for the Living Collections is now in its second year and it is already providing greenhouse staff with exciting and challenging opportunities to germinate seeds, root cuttings, and grow-on wild-collected species that are new to the Arboretum as well as previously attempted taxa. The Campaign has reinforced the importance of horticultural research and reasserts that propagation is very much center stage, even as we near our 2022 sesquicentennial. As autumn is fast approaching and new collecting expeditions will soon start, we are once again awaiting the propagules that will be beginning their journey through the production system. We can only imagine that this is how Dawson felt during Wilson's 1907 to 1909 expedition to China, eager to receive the 2,262 seed collections and 1,473 collections of live plants or cuttings that resulted from the trip. References Dosmann, M. 2015. The History of Minimum Temperatures at the Arnold Arboretum: Variations in Time and Space. Arnoldia 72(4): 2?11. Dosmann, M. and K. Port. 2016. The Art and Act of Acquisition. Arnoldia 73(4): 2?17. Friedman, W. E., M. S. Dosmann, T. M. Boland, D. E. Boufford, M. J. Donoghue, A. Gapinski, L. Hufford, P. W. Meyer, and D. H. Pfister. 2016. Developing an Exemplary Collection: A Vision for the Next Century at the Arnold Arboretum of Harvard University. Arnoldia 73(3): 2?18. Geary, S. C. and B. J. Hutchinson. 1980. Mr. Dawson, Plantsman. Arnoldia 40(2): 51?75. Howard, R. 1962. The Charles Stratton Dana Greenhouses of the Arnold Arboretum. Arnoldia 22(5?6): 33?47. Wilson, E. H. 1930. Bulletin of Popular Information. The Arnold Arboretum of Harvard University (series 3, vol 4, no 10): 37?40. Wilson, E. H. 1906. Some New Chinese Plants. Bulletin of Miscellaneous Information. Royal Botanic Gardens, Kew. 1906(5): 147?163. Wyman, D. 1960. Plants of Possible Merit? Arnoldia. 20(2): 9?16. Tiffany Enzenbacher is Manager of Plant Production at the Arnold Arboretum. Plant Propagator John H. Alexander III recently retired from the Arboretum after 40 years of service. "},{"has_event_date":0,"type":"arnoldia","title":"Unlocking Ancient Environmental Change with the Help of Living Trees","article_sequence":2,"start_page":14,"end_page":22,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25608","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170816f.jpg","volume":74,"issue_number":1,"year":2016,"series":null,"season":null,"authors":"Marston, John M.","article_content":"Unlocking Ancient Environmental Change with the Help of Living Trees John M. Marston W ith both human societies and ecosystems worldwide now facing ongoing, and even accelerating, environmental change, both scholars and policy makers are increasingly concerned with predicting the future implications of climate change. Where will our coastlines, tree lines, and urban boundaries lie in 50 or 100 years? How will changes in the seasonality and intensity of precipitation, frosts, and heat waves affect the plants and animals on which we rely for food? And, most important, what are the consequences for us? One avenue for understanding human responses to dramatic environmental and climatic change is to look to the past when societies faced similar periods of rapid change. Paleoclimatologists and paleoecologists have developed numerous methods to identify ancient environmental change, creating rich records from glacial ice at the poles and on mountaintops, as well as cores drilled deep into seabeds and lakes that preserve hundreds or thousands of years of annually deposited sediments. Archaeologists who study the deep history of human-environmental relationships draw on these datasets, as well as archaeological records of social and economic change, to explore human adaptation to environmental change in the past. A variety of archaeological finds are useful in identifying climatic change, from mammal and fish bones to microscopic starch grains found on tools used in plant food processing. One material commonly found in archaeological sites from many different periods of the human past, nearly worldwide, is wood charcoal. Incompletely burned wood from fireplaces, ovens, kilns, and accidently (or deliberately) burned buildings becomes inorganic charcoal, which is resistant to degradation from soil microbes and fungi and thus can survive for thousands of years within the soil. It is frequently possible to identify the type of tree that produced these charcoal remains and thus reconstruct patterns of wood use and forest change, both as a result of climatic change and deliberate or inadvertent human reconfiguration of woodlands. Scholars have developed methods for systematically recovering, identifying, and interpreting these remains to identify patterns of climate and environmental change in the past. Recently, Boston University and the Arnold Arboretum have begun a partnership to draw on the vast living collections of the Arboretum to improve the resolution of archaeological charcoal studies in the Environmental Archaeology Laboratory in the Department of Archaeology at Boston University. In this article, I describe how archaeologists study charcoal from archaeological sites and use it to reconstruct the human role in environmental change, highlighting how resources of the Arnold Arboretum enhance our teaching and research mission at Boston University. Recovering and Identifying Archaeological Plant Remains Wood charcoal fragments from archaeological sites have been studied since the 1940s to address multiple questions about human wood use in the past. The first step in archaeological charcoal analysis is systematic recovery of charcoal remains from archaeological sites. Although not a universal practice, the recovery of plant remains is increasingly ubiquitous among archaeologists worldwide, even in remote areas of developing countries. We recover soil samples, generally 10 to 20 liters (2.6 to 5.3 gallons) in volume (equivalent to one or two buckets full), from every archaeological level and distinct feature (e.g., a pit or a hearth) identified during excavation. Archaeologists most commonly use a water flotation method to recover charred plant ALL IMAGES BY THE AUTHOR UNLESS OTHERWISE INDICATED Ancient Environmental Change 15 The author operating a flotation tank on site in Turkey, and charred plant remains floating to the surface within the tank. remains, including wood charcoal as well as carbonized seeds and other plant structures, from soil samples. Although flotation can be accomplished using only a pair of buckets and a fine mesh strainer, more common are systems that pump large volumes of water to process even large samples quickly. Clean water is pumped into the tank of the machine where the soil sample is held in a plastic window screen mesh. The water dissolves the soil, freeing carbonized plant remains, which float, and rinsing away sediment in the dirty effluent that is released from the bottom of the tank. Heavy components of the soil, including bone and pottery fragments as well as occasional heavy pieces of charcoal, are caught in the window screen and later dried and analyzed. The floating, or light, fraction consists of wood charcoal and carbonized plant remains, but also soil components lighter than water, including tiny roots and fine clay particles. The light fraction is allowed to overflow into a very fine polyester mesh, with holes less than 0.1 millimeter (0.004 inch) to catch even the smallest seeds. This fraction is then carefully air dried and brought to the laboratory for identification and analysis. We then pour the light fraction through a series of nested sieves, creating several size classes of material that can be sorted differently. In general, only wood charcoal fragments larger than 2 millimeters (0.08 inch) are analyzed, as smaller fragments are unlikely to be identifiable. Systematically sorting each size class under low-power stereomicroscopes, we remove each type of plant remain for subsequent identification and measurement, with wood charcoal, carbonized seeds and seed fragments, and nutshell distinguished and separated. Wood charcoal fragments are then weighed in aggregate and a representative number of those fragments are identified. The identification of wood charcoal can be challenging because fragments are often small and may be distorted by burning and subsequent deterioration in the soil. Fortunately, different species of woody plants vary considerably in their cellular anatomy, which allows wood (even charcoal) to be identified to varying levels of specificity depending on the wood 16 Arnoldia 74\/1 ? August 2016 (Left) Diagram of pine wood, showing three planes of structure (image from Plant Anatomy by William Chase Stevens, 1916, Philadelphia: P. Blakiston. Courtesy of Florida Center for Instructional Technology, http:\/\/etc. usf.edu\/clipart\/). (Right) Scanning electron micrograph of Turkey oak (Quercus cerris) wood from the Environmental Archaeology Laboratory collection. (Left) Sugar maple (Acer saccharum) wood, in transverse section (scale bar 500 ?m = 0.5 mm); growth ring boundary is marked with red line. (Right) Black pine (Pinus nigra) wood, in transverse section (scale bar 500 ?m = 0.5 mm); growth ring boundaries are marked with red lines. type. Wood can be viewed from three planes, each of which presents a distinct set of anatomical structures for identification. All three are necessary for detailed identification, but the transverse, or cross section, is the most useful for charcoal identification and can be examined with a stereomicroscope at 20 to 100? magnification. Distinguishing hardwoods (angio- sperms) and softwoods (gymnosperms) can be easily accomplished using just low-power magnification of the transverse section; many families within these large categories can also be distinguished based solely on the transverse section. Using a combination of basic reflected light microscopy, high-power incident light microscopy, and electron microscopy, we cata- Ancient Environmental Change 17 log features of archaeological wood fragments and assign them tentative identifications based on their anatomy. Confirmation of these identifications, however, typically requires a comprehensive comparative collection of modern wood taken from properly identified and fully vouchered trees. Assembling such a comparative collection has been an ongoing effort of the Environmental Archaeology Laboratory and is the origin of our collaboration with the Arnold Arboretum. Using the Arboretum as a Research Collection The Arnold Arboretum offers a tremendous opportunity to collect wood from a wide variety of temperate tree species from the Americas, Europe, and Asia. Each tree is properly identified and labeled, and considerable information regarding its life history is recorded in the Arboretum's living collections database. For our partnership, since most woody plants are identifiable at the genus level, we preferentially collect wood from species native to the areas in which members of the Environmental Archaeology Laboratory work (mainly southern Europe, the Middle East, East Asia, and northeastern North America). When the most relevant species are not available, we choose other species of those genera in order to obtain the most similar comparative specimens possible. Wood anatomy can vary based on the diameter and age of the branch collected, between branch and trunk wood, and because of unique growth conditions such as bending or disease. As a result, we attempt to collect wood from multiple parts of a tree when possible. The Arboretum facilitates our collection by allowing us to gather dead branches that have fallen from trees as well as gathering samples from trees that are trimmed or cut down during the course of routine tree maintenance activities. Members of the Environmental Archaeology Laboratory compiled a \"wish list\" of trees in the living collections that Arboretum arborists can refer to when tree work is done. The arborists then collect specimens from trees of specific interest to us. We periodically stop by the Arboretum to collect these wood samples for further processing at Boston University. Back in the Environmental Archaeology Laboratory, we interface with the Arboretum's database and use the Arboretum Explorer website (http:\/\/arboretum.harvard.edu\/explorer\/) to gather information about trees that have been sampled. We record much of that information into the Environmental Archaeology Laboratory Collections Database, which is also searchable online (http:\/\/sites.bu.edu\/ealab\/ collections\/database\/). The wood sample is then divided between a wood specimen and a specimen to be converted into charcoal. Experimental carbonization of comparative wood samples is critical for two reasons. First, carbonization can modify the structure of the wood in predictable ways, leading perhaps to certain patterns of cracks that can be diagnostic when examining archaeological wood charcoal. Second, charcoal can be easily broken to expose any of the three planes, facilitating rapid examination, while wood needs to be cut with an ultrathin blade so as not to crush the exposed cell walls, requiring additional equipment and time to prepare comparative slides. We carbonize wood using a muffle furnace capable of reaching temperatures of 1000?C (1832?F), although we typically carbonize wood around 400?C (752?F) to maximize speed of carbonization without incinerating the wood. It is critical that wood heat in an oxygen-poor reducing atmosphere because that promotes charcoal formation, while an abundance of oxygen would lead to ashing and destruction of the sample. We carefully wrap samples twice in heavy-duty aluminum foil to minimize contact with oxygen and pack them tightly in the muffle furnace. At 400?C, wood carbonizes in 10 to 40 minutes, depending on the thickness of the pieces. Finally, both charcoal and wood specimens are stored in labeled boxes within a specialized shelving system in the lab. The boxes include basic information on the wood and its location of origin, together with an identifier code that corresponds to its record in our database. A future project for the laboratory is to take microscopic images of the wood anatomy of all woods in the collection and to make them available online, both through the laboratory website and as a contribution to Inside Wood 18 Arnoldia 74\/1 ? August 2016 Clockwise from top left: Collected wood specimens to be accessioned into comparative collection. Larger pieces were provided by the Arnold Arboretum. Preparation of wood for experimental carbonization: sawing a sample for carbonization; packing the muffle furnace; a fully carbonized specimen, just out of the furnace. Samples housed and labeled for comparative collection. Ancient Environmental Change 19 Once it is possible to identify wood fragments reliably, we work to identify a statistically robust subsample of all wood charcoal fragments present in our archaeological samples. Recording both count and weight of these fragments, we are able to create diagrams that represent change in the prevalence and context of use for woods over time. For example, in my ongoing research at the ancient city of Gordion, in central Turkey, which was inhabited from the Early Bronze Age (3000 to 2000 BC) through the medieval period (fourteenth century AD), I was able to document changes in wood use practices and forest ecology over a span of 3,000 years. Gordion became a large city around 800 BC as the capital of the Phrygian kingdom, which grew from Gordion to control most of central Turkey. At that time the Phrygians began to construct monumental temples, massive city walls, and huge earthen burial mounds (the largest over 170 feet [52 meters] in height) containing royal burials inside elaborate wooden structures, including the oldest standing wooden building in the world. This amazing structure was fashioned from juniper (Juniperus spp.) wood, which was widely used within the city in roofing large public buildings. Juniper is a slow-growing tree, however, and the inhabitants of Gordion appear to have quickly exhausted their supply of easily cut large juniper trees. In later periods of occupation, charcoal samples from burned buildings indicate that oak (Quercus spp.) and pine (Pinus Wood samples from this yellow birch (Betula alleghaniensis, accession 629-83-F) were carbonized for the Boston University Environmental Archaeology Laboratory charcoal collection. KYLE PORT Reconstructing Past Woodland Ecology and Wood Use, with Implications for the Future KYLE PORT (http:\/\/insidewood.lib.ncsu.edu), a free, public, wood anatomy database created at North Carolina State University. Although extensive comparative collections of wood samples are preserved at other arboreta and herbaria worldwide, very few of these have been digitized to make them publically accessible. Because our collection includes specimens from many countries of the Middle East and Central Asia, as well as specimens from several arboreta in the United States, we aim to publicize our records as widely as possible as a research tool for archaeologists worldwide. Wood samples from this hybrid tuliptree (Liriodendron tulipifera ? chinense, accession 584-81-A) growing near the Arboretum's Hunnewell Visitor Center were provided to the Boston University Environmental Archaeology Laboratory. Using the Arboretum as a Teaching Collection Every summer I teach a weeklong intensive workshop on wood anatomy and wood charcoal identification for archaeologists. Participants come to Boston from universities nationwide, from Santa Barbara to Chapel Hill, and a few even join us from across the river in Cambridge. Participants are mainly doctoral students, but we have a number of faculty participants and even an occasional undergraduate. During the week we cover wood anatomy from initial concepts (e.g., the three planes of wood) to advanced structural variation (e.g., ray cell margin shape in gymnosperms). We also read and discuss a number of articles that illustrate best practices for sampling and recovery of wood charcoal from archaeological sediments, methods for quantifying and presenting results, and the challenges of changes brought about by both the initial burning of wood and its preservation in soils for hundreds or thousands of years. Students spend the last few days analyzing their own wood charcoal assemblages and learning how to identify the woods common to their areas of expertise, which have ranged from southwest China to Jordan, the California Channel Islands, the Yucatan, and the Andes. One highlight of the week is our field trip to the Arnold Arboretum. The group receives an orientation and tour led by Michael Dosmann, Curator of Living Collections. During the tour, workshop participants learn about the unique collections of the Arnold Arboretum and about the life history of particular trees on the property. Following an orientation to the Arboretum Explorer web application, participants are able to use their smartphones to find particular trees of interest to them and spend the next two hours visiting those trees. We collect a few samples of dead wood from the ground under selected trees to bring back to the laboratory, where we then experimentally carbonize wood samples and study their microscopic structure. Participants then split their newly collected specimens, with a portion joining the permanent collection of the Environmental Archaeology Laboratory and the rest returning home with each participant. As a result, every participant returns to their home lab with the beginning of a personal comparative charcoal collection and the experience needed to expand their collection through fieldwork and collaborative ventures with local botanical gardens and arboreta. Participants in the 2014 wood charcoal workshop analyzing samples in the Boston University Environmental Archaeology Laboratory. Ancient Environmental Change 21 Clockwise from top: The funerary chamber within the largest burial mound at Gordion, dated to 743?741 BC, showing the outer casing of roughly finished juniper logs. Juniper logs used as support ties within a stone wall at Gordion. A Greek juniper (Juniperus excelsa) of a size similar to that of the logs in the funerary chamber; trees of this size are rare in the landscape around Gordion today. 22 Arnoldia 74\/1 ? August 2016 spp.) were the primary woods used in construction, both of which have the advantage of being fast-growing trees that often take over in sites where older juniper trees have been cut. Oak and pine, however, have inferior strength and rot resistance compared to juniper. Archaeological wood charcoal assemblages show a dramatic human impact on the landscape that led to considerable forest reorganization during the early history of the city. Later inhabitants of the region had to contend with a different landscape, and different availability of natural resources, than their ancestors. Examples such as the case of Gordion parallel more recent human history, both in central Turkey and worldwide, in which human activity transforms a landscape for future inhabitants. When viewed from the perspective of later populations, we term these impacts \"legacy effects,\" and the implications of such changes are many. It has been argued by several scholars, including Jared Diamond, that the deforestation of Easter Island pushed its ecosystem beyond a tipping point that led to severely reduced resources and impoverishment of the isolated inhabitants. In contrast, legacy effects may also have been deliberate outcomes, designed to boost productivity and resource availability. The use of fire to maintain prairie habitats in the American Great Plains prior to European contact is an example of such \"niche construction,\" in which people modify their environment to boost productivity of desired resources to suit their cultural needs. Archaeologists have explored these environmental histories using wood charcoal analysis, and continue to search for a deeper understanding of not only when and how, but also why human groups manipulate their landscape in specific ways. These detailed studies offer cases of environmental disaster and social collapse, but also resilience and survival in even the most uninviting landscapes. As contemporary society faces environmental change on an unprecedented scale, archaeologists offer both cautionary and inspiring stories of humanenvironmental relationships that provide novel, proven effective tools for continued survival in a changing world. Additional Reading These include sources that outline the practice of archaeological wood charcoal analysis (Asouti and Austin 2005, Marston 2009); wood anatomy and identification (Panshin and de Zeeuw 1970, Schweingruber 1990, Schweingruber et al. 2006); frameworks for studying human-environmental interactions (Cumming et al. 2006, Marston 2015, Redman 1999, Smith 2007); and more about our team's recent work at Gordion (Marston in press, Miller 2010, Rose 2012). Asouti, E. and P. Austin. 2005. Reconstructing woodland vegetation and its exploitation by past societies, based on the analysis and interpretation of archaeological wood charcoal macro-fossils. Environmental Archaeology 10: 1?18. Cumming, G. S., D. H. M. Cumming, and C. L. Redman. 2006. Scale mismatches in social-ecological systems: causes, consequences, and solutions. Ecology and Society 11: 14. Marston, J. M. 2009. Modeling wood acquisition strategies from archaeological charcoal remains. Journal of Archaeological Science 36: 2192?2200. Marston, J. M. 2015. Modeling resilience and sustainability in ancient agricultural systems. Journal of Ethnobiology 35: 585?605. Marston, J. M. (In press). Agricultural Sustainability and Environmental Change at Ancient Gordion. Philadelphia: University of Pennsylvania Museum Press. Miller, N. F. 2010. Botanical Aspects of Environment and Economy at Gordion, Turkey. Philadelphia: University of Pennsylvania Museum Press. Panshin, A. J. and C. de Zeeuw. 1970. Textbook of Wood Technology. New York: McGraw Hill. Redman, C. L. 1999. Human Impact on Ancient Environ ments. Tucson: University of Arizona Press. Rose, C. B. (editor). 2012. The Archaeology of Phrygian Gordion, Royal City of Midas. Philadelphia: University of Pennsylvania Museum Press. Schweingruber, F. H. 1990. Anatomy of European Woods. Stuttgart: Haupt. Schweingruber, F. H., A. B?rner, and E. D. Schulze. 2006. Atlas of Woody Plant Stems: Evolution, Structure, and Environmental Modifications. Berlin: Springer. Smith, B. D. 2007. The ultimate ecosystem engineers. Science 315: 1797. John M. Marston is Assistant Professor in the Departments of Archaeology and Anthropology at Boston University. "},{"has_event_date":0,"type":"arnoldia","title":"Cork: Structure, Properties, Applications","article_sequence":3,"start_page":23,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25606","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170bb26.jpg","volume":74,"issue_number":1,"year":2016,"series":null,"season":null,"authors":"Gibson, Lorna J.","article_content":"Cork: Structure, Properties, Applications Lorna J. Gibson BIODIVERSITY HERITAGE LIBRARY E ver since people have cared about wine, they have cared about cork to keep it sealed in bottles. \"Corticum abstrictum pice demovebit amphorae ...\" (Pull the cork, set in pitch, from the bottle) sang the Roman poet Horace in 27 B.C., to celebrate the anniversary of his miraculous escape from death from a falling tree. In Roman times, corks used to seal bottles were covered in pitch; it was not until the 1600s that a method for stoppering bottles with clean corks was perfected by Benedictine monks at Hautvillers in France. Cork's elasticity, impermeability, and chemical stability means that it seals the bottle without contaminating the wine, even when it must mature for many years. The Romans also used cork for the soles of shoes and for floats for fishing nets. According to Plutarch (A.D. 100), when Rome was besieged by the Gauls in 400 B.C., messengers crossing the Tiber clung to cork for buoyancy. Cork is the bark of the cork oak, Quercus suber, which grows in Mediterranean climates. Pliny, in his Natural History (A.D. 77), describes it: \"The cork-oak is a small tree, and its acorns are bad in quality and few in number; its only useful product is its bark which is extremely thick and which, when cut, grows again.\" All trees have a thin layer of cork in their bark; Quercus suber is unusual in that, at maturity, the cork forms a layer many centimeters thick around the trunk of the tree. The cell walls of cork are covered with thin layers of unsaturated fatty Robert Hooke's book Micrographia amazed readers with its detailed drawings such as this one of cork showing the roughly rectangular cell shape in one plane and the roughly circular cell shape in the perpendicular plane. The lower drawing is of sensitive plant (Mimosa pudica), whose touch-induced leaf movement Hooke studied. For more images and insight on Micrographia from this article's author, please view this YouTube video: https:\/\/www. youtube.com\/watch?v=zFfVtziLhg4 24 Arnoldia 74\/1 ? August 2016 COURTESY OF AMORIM, COPYRIGHT APCOR (PORTUGUESE CORK ASSOCIATION) Cork is harvested from managed cork oak (Quercus suber) forests such as this one in Portugal. acid (suberin) and waxes, which make them impervious to air and water, and resistant to attack by many acids. Cork Under the Microscope shape in the perpendicular plane. Hooke noted that the cell walls were arranged \"as those thin films of Wax in a Honey-comb.\" Modern scanning electron micrographs of cork show additional detail. In the plane in which the cells look rectangular, we see that the cell walls are wavy, rather than straight, and in the perpendicular plane, the cells are roughly hexagonal prisms, with the waviness in the cell walls along the length of the prism axis. The dimensions on the unit cell are microns, or micrometers (?m); for comparison, a human hair is roughly 50 microns in diameter. Cork occupies a special place in the history of microscopy and of plant anatomy. When English scientist Robert Hooke perfected his microscope, around 1660, one of the first materials he examined was cork. What he saw led him to identify the basic unit of plant and biological structure, which he called the \"cell\" (from cella, Latin for small chamber). His book, Micrographia, published in 1665, records his observations, including the comment that, \"I no sooner descern'd these (which were indeed the first microscopical pores I ever saw, and perhaps, that were ever seen, for I had not met with any Writer or Person that had made any mention of them before this) but me thought I had with the discovery of them, presently hinted to me the true and intelligible reason of all the Phenomena of Cork.\" Hooke's detailed drawings of cork show the Scanning electron micrographs of cork cells in the same two perpendicular roughly rectangular cell shape in one planes as in Hooke's drawings, showing the corrugations in the cell walls plane and the roughly circular cell (from Gibson et al., 1981). Cork Tree vs Cork Oak LESLIE J. MEHRHOFF, UCONN, BUGWOOD.ORG The Arboretum's cork tree (Phellodendron spp.) collection lies south of the Hunnewell Visitor Center along Meadow Road, as seen in the photo below. There are 18 Phellodendron specimens comprising 5 taxa, all native to Asia, in the collection. While the bark of cork trees has a similar compliant feel as that of the true cork oak (Quercus suber), it is not used as a source of cork. These scanning electron micrographs of two perpendicular planes in Phellodendron bark show more irregular cells compared with those of Quercus suber (seen on page 24). Cork oak is only cold hardy through USDA hardiness zone 8 (average annual minimum temperature 10 to 20?F [-12.2 to -6.7?C]) so there are no specimens at the Arboretum. 26 Arnoldia 74\/1 ? August 2016 Sheets of cork oak bark rest in front of the tree they were harvested from. KESSLER AND SONS MUSIC Cork is roughly 15% solid and the rest is air. Its density is typically about 15% that of water: its low density, combined with the closed cells that do not allow water to enter, gives cork its great buoyancy. The low volume fraction of solid, along with the relatively compliant cell wall material, gives rise to its compressibility. The waviness or corrugations in the cell walls of cork leads to an unusual behavior: if pulled along the prism axis, the corrugations in the cell walls straighten out, with little change in the transverse dimension (like the bellows of an accordion unfolding). In contrast, if you pull on most materials they get narrower in the transverse direction (think of pulling on a rubber band, for example). And if a cube of rubber is compressed some amount in one direction, it will expand out sideways by nearly half that amount in each of the other two transverse directions. When compressed along the prism axis, the corrugations in cork's cell walls simply fold up, again producing no change in the transverse dimension. It is this property, along with the compressibility of cork, that makes it easy to insert cork into a bottle and gives a good seal against the glass neck of the bottle. Cork makes good gaskets for the same reason that it makes good bungs for bottles: it is compressible, accommodating deformation, and its closed cells are impervious to liquids. Thin sheets of cork are used, for instance, as gaskets between sections of woodwind instruments. The sheet of cork is always cut with the prism axis normal to its plane, so that when the two sections of the instrument are mated, the cork does not expand around the circumference of the section and will not wrinkle. Cork makes an admirable flooring material because it is comfortable to walk on (thanks to its compressibility), it holds warmth, and it doesn't become slippery, even when wet. Cork holds warmth because it transfers heat poorly. In porous, cellular solids such as cork, heat transfer occurs by conduction (through the solid or gas), by convection (as gas on the warmer side of a cell rises and that on the cooler side falls, setting up convection currents), or by radiation. Gases have lower thermal conductivities than solids (by a factor of up to a thousand) so COURTESY OF AMORIM How Cork Works Cork gaskets are used at the tenon joints of clarinet sections. the high volume fraction of air within the cells reduces heat transfer by conduction through cork. Convection currents, carrying heat from one side of a cell to the other, are suppressed for cell sizes less than about 1 millimeter (for small cell sizes, the buoyancy force associated with hot air rising is counteracted by drag of the air against the walls of the cells). And heat flow by radiation also depends on cell size--the smaller the cells, the more times the heat has to be absorbed and reradiated, reducing the rate of heat flow. So the high volume fraction of air in cork and its small cells contribute to its ability to hold warmth. Cork 27 unloaded loaded in tension Micrographs showing (left) cork cells unloaded and (right) the progressive straightening of cell walls as cork is pulled along the prism axis (from Gibson et al., 1981). A pin pushed into cork results in a narrow band of crushed cells next to the pin (left) but little deformation of the cork beyond that (right) (from Gibson et al., 1981). Friction between a shoe and a cork floor has two origins. One is adhesion, in which atomic bonds form between the two contacting surfaces and work must be done to break them. Between a shoe and a tiled or stone floor, this is the only source of friction, and since it is a surface effect, it is completely destroyed by a film of water or soap, making the floor slippery. The other source of friction is due to energy losses associated with loading and unloading the floor (as a step is taken, for instance). In some materials, such as stone, these energy losses are small, but in cork, the energy losses are significant (it is said to have a high loss coefficient). Since the energy losses occur within the cork, and are not a surface effect, cork floors do not become slippery even when wet or soapy. Cork is widely used for bulletin boards. When a pin is stuck into cork, the deformation is very localized around the pin. A narrow band of cork cells, occupying a thickness of only about a quarter of the diameter of the pin, collapses, crushing those cells nearly completely, to accommodate the diameter of the pin. The deformation in the cells beyond this highly deformed band is negligible in comparison. For this reason, the force needed to push the pin into a cork bulletin board is small. And cork recovers most of the deformation when it is unloaded, so that the hole nearly closes up after the pin is removed. The cellular structure of cork is unique. It gives rise to a remarkable combination of properties that are exploited in everything from bottle stoppers and gaskets to the soles of shoes, flooring, and bulletin boards. Acknowledgements This article is based on the paper, The structure and mechanics of cork, co-authored with Ken Easterling and Mike Ashby, referenced below; it is a pleasure to acknowledge their contributions. Micrographs on pages 24 and 27 are from that paper. References Gibson, L. J. , K. E. Easterling, and M. F Ashby. 1981. Structure and mechanics of cork. Proceedings of the Royal Society, A377, 99?117. Hooke, R. 1665. Micrographia, Tab XI. London: Royal Society. Horace, Q. circa 27 BC. Odes, book III, ode 8, line 10. Pliny, C. 77 AD. Natural History, vol. 16, section 34. Plutarch. 100 AD. Life of Camillus. Parallel Lives, vol. 2, ch. 25, p. 154. Lorna J. Gibson is the Matoula S. Salapatas Professor of Materials Science and Engineering at the Massachusetts Institute of Technology. "},{"has_event_date":0,"type":"arnoldia","title":"Ulmus thomasii: The Hard Elm That's Hard to Find","article_sequence":4,"start_page":28,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25607","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170bb6b.jpg","volume":74,"issue_number":1,"year":2016,"series":null,"season":null,"authors":"Pruka, Brian","article_content":"Ulmus thomasii: The Hard Elm That's Hard to Find Brian Pruka I f you hike Beech Path up the steep slope from Valley Road and continue to the point where a footpath branches off to the right, you will find a slender, stately tree next to the trail. It is a rock elm, Ulmus thomasii, one of only three elms native to northeastern North America. This particular specimen, accession number 444-88-A, is the only rock elm currently in the Arnold Arboretum's living collections. It is well worth seeing. Rock elm was originally named Ulmus rac emosa in 1831 by its discoverer, American civil engineer David Thomas of New York. It was renamed Ulmus thomasii in 1902 by Arboretum director Charles Sprague Sargent when he determined that another elm already had the name Ulmus racemosa. Rock elm is most common in the northeastern and north-central states, with the core of its range stretching from north-central Wisconsin to southern Michigan and southern Ontario. Populations exist as far south as Tennessee, but it is primarily a cold-weather tree, not often found in regions warmer than USDA Plant Hardiness Zone 5 (average annual minimum temperature -10 to -20?F [-23.3 to -28.9?C]). Rock elm is rarely encountered in New England, likely because it has a strong preference for limestone substrates, which are not common here. Back in the 1910s to 1930s there were as many as twelve rock elms growing on the Arboretum grounds, all procured from well known plant nurseries of the era. Most of these trees eventually died of Dutch elm disease (DED), a devastating fungal vascular wilt. Four succumbed to Boston's first big DED epidemic in 1946. Two died of DED in 1987, another three in 1989. Specimen 17925-B was recorded as being in \"excellent health\" on May 5, 1989. It was cut down 75 days later, on July 19, dead from DED. Our current living specimen was propagated in 1988 as a cutting from a then 102-year-old tree (accession 17926-A) that was planted at the Arboretum in 1886. One of the best traits for identifying a rock elm--not often listed in identification books-- is its form. The species is typically tall and slender, with a single bole that gets remarkably tall before it splits into a narrow crown. Rock elms growing in crowded forest situations also usually have small corky branches that droop downward from the middle third of the main bole. The Arboretum's specimen has grown out in the open all its life and does not have drooping branches at mid-bole, though it does have a strikingly straight main trunk and currently measures 44.24 feet (13.48 meters) tall with a dbh (diameter at breast height) of 18.31 inches (46.5 centimeters). Rock elm leaves can look much like American elm leaves. Tree identification books generally list three identifier traits for rock elm: branches with 3 to 5 irregular corky wings; inflorescences of 7 to 13 flowers arranged in a long, pendulous raceme; and fruits (samaras) covered with tiny hairs and an inflated paper wing that is not distinct from the seed case. Unfortunately those unique traits are not always present. Some rock elms, including our specimen, lack corky twigs. Rock elms don't reproduce until about age 20, don't produce full seed crops until age 45, and produce bountiful seed crops only once every 3 to 4 years. Seeds drop from the tree as soon as they ripen, so from May to February there are no reproductive structures to aid identification. The timber of rock elm is especially prized for its hardness. It has interlaced fibers that make it almost impossible to split, yet easy to bend. It is especially durable underwater. In past centuries, much rock elm was cut and shipped to Great Britain to build wooden battleships. Rock elm is also highly regarded for its beautiful gold fall foliage color, so consider a hike up Beech Path this autumn. A tall, handsome native elm is awaiting you. Brian Pruka is a 2016 Isabella Welles Hunnewell Intern at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23450","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd1708928.jpg","title":"2016-74-1","volume":74,"issue_number":1,"year":2016,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Art and Act of Acquisition","article_sequence":1,"start_page":2,"end_page":17,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25604","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170b728.jpg","volume":73,"issue_number":4,"year":2016,"series":null,"season":null,"authors":"Port, Kyle; Dosmann, Michael S.","article_content":"CAMPAIGN FOR THE LIVING COLLECTIONS The Art and Act of Acquisition Michael S. Dosmann and Kyle Port The Campaign for the Living Collections is well under way at the Arboretum. In the last issue we presented the document that guides the Campaign, and in this issue we get to the action-- how does the Arboretum curatorial staff prepare for and carry out plant collecting expeditions? Curator of Living Collections Michael Dosmann and Manager of Plant Records Kyle Port provide insight and share photographs from their recent trips. P lant exploration combines a love of plants with adventure. Over its nearly 145-year history, the Arnold Arboretum has harnessed these passions by leading or supporting more than 150 plant collecting events across 70 countries. As knowledge of the plant kingdom has evolved, so have the Arboretum's living collections, placing even greater demand on deliberate and strategic collection planning. As described in the previous issue of Arnoldia (Friedman et al. 2016), the new 10-year Campaign for the Living Collections articulates a number of broad goals that, when met, will preserve the collections' singular legacy and advance it well into the future. For example, there is a call to strengthen the species representation within genera such as Viburnum and Taxus that are useful to the study of biogeography. As a means of broadening the number of genera in the collections, several marginally hardy taxa like Daphniphyllum macropodum and Nothofagus dombeyi have been identified as species worth trying to grow here. And, because of the great threat of extinction, numerous conservation-status species are highlighted before they disappear from the wild. The Arboretum will meet these collections goals through the acquisition of nearly 400 target taxa, or desiderata, with each fulfilling at least one (and typically several) goals. For many of the taxa on the list, the Arboretum needs several unique acquisitions (e.g., from multiple locations), so what is initially a list of 395 blossoms into a vibrant garden of 720 actual targets. Each of these targets will require its own acquisition plan and approach. A few might be purchased from nurseries, some may be acquired from cooperative institutions and repositories, while others will be sought out and obtained through the Arboretum's network of colleagues. However, the majority will be obtained on specific plant expeditions in which an Arboretum staff member leads or participates. With the vision and goals in place, a new generation of explorers, horticulturists, and other Arboretum friends and associates are rallying to collect from the temperate flora and cultivate these plants in the Arboretum. Few endeavors are as rewarding and exciting as seeing plants in their wild habitats, collecting seeds or other propagules, and then bringing them back home to cultivate. As botanical garden professionals, we also value the role plant exploration plays in other aspects of collections manage- Smoke from dynamic forest fires cloaked the Alpine Lakes Wilderness in the central Cascade Mountains of Washington as Kyle Port's flight descended into the region at the beginning of his plant collecting trip to northern Idaho last fall. ment. The important pre-trip planning and posttrip follow-up lead not only to better executed expeditions, but to more individual (and thus institutional) knowledge that can be applied to the long-term stewardship of the species collected. In this article, we shed some light on a number of aspects of plant collecting based on some recent trips and our shared experiences. It is not our intention to be encyclopedic on the subject, however, as many resources exist that cover in greater depth. (For more details on the philosophy, mechanics, and best practices of plant exploration see Ault (2000), ENSCONET (2009), and the special 2010 issue of Arnoldia (Volume 68, Number 2) commemorating the 20th-Anniversary of the North America-China Plant Exploration Consortium [NACPEC]. For the majority of the expeditions that will be conducted by the Arboretum over the next decade, the various desiderata will dictate where (and how) we collect them. Simply put, as the individual targets are marked on maps, regions that contain the most \"dots\" become obvious sites for exploration. A region with just a single species might be best left to a contrac- tor or other means. With respect to the Campaign, Living Collections Fellow Robert Dowell has been tasked with conducting species audits to identify these ideal collecting localities. For the inaugural expeditions that marked the Campaign's launch in the autumn of 2015, we (the authors) preselected regions based on our prior knowledge of the areas and confidence that they would yield quality material listed as desiderata. Kyle collected in northern Idaho from August 24 to September 4, 2015; Michael participated in a NACPEC expedition in central China focused on paperbark maple (Acer griseum) from September 1 to 18, and then struck out with another set of colleagues in northern Sichuan from September 18 to 29. BE PREPARED In many respects, a trip's success--both shortand long-term--is proportional to the amount of planning. Once a general region for an expedition has been selected, the trip participants hone their knowledge about the target species' biology, including their identification, phenology (e.g., what time of year mature seeds are KYLE PORT 4 Arnoldia 73\/4 NANCY ROSE Acquisition 5 ANTHONY S. AIELLO Local collaborators are invaluable on expeditions. (Left) Kang Wang, from the Beijing Botanical Garden, has acted as liaison for a number of expeditions including the 2015 NACPEC trip that concentrated on paperbark maple (Acer griseum). Michael Dosmann, Kang Wang, and Kris Bachtell (left to right) sort and organize Acer griseum leaves that will later be subject to DNA extraction. (Right) Paul Warnick, from the University of Idaho Arboretum and Botanical Garden, collaborated with Kyle Port on the 2015 North Idaho Expedition. He's seen here with a collection of devil's club (Oplopanax horridus) in the Selkirk Mountains, Kaniksu National Forest, Idaho. What's a Collection? The word \"collection\" can refer to a group of living plants, like the Lilac (Syringa) Collection at the Arboretum. But on plant collecting expeditions, \"collection\" is also the term used for the products of each unique act of collecting. For example, if both seeds and herbarium vouchers are collected from a tree, a single collection number is assigned to both since they came from the same plant. When the seeds are distributed to other institutions, each institution will assign an accession number from its own system, but the original collection number will be included in the accession information so the original source can always be traced. MICHAEL S. DOSMANN Anthony S. Aiello measures out seeds of Ostrya japonica, collection number 24 from the NACPEC 2010 trip, prior to packaging and shipping them back to the United States. KYLE PORT 6 Arnoldia 73\/4 KYLE PORT Acquisition 7 KYLE PORT (Left) David Port (Kyle's father), raised in the Idaho hamlet of Troy, played many roles over the course of the 2015 North Idaho Expedition, including lugging the pole pruners to the edge of Lake Pend Oreille and to the high elevations around Roman Nose Lakes. (Right) Pole pruners also came in handy during the 2014 Ozarks expedition, where they were used to collect these fruit capsules of red buckeye (Aesculus pavia) in Arkansas. forms, printed on waterproof paper, is best for all weather situations. The recorder must have a keen attention to detail, be thorough, and, of course, have excellent plant knowledge. Memories always fail, so it's important to capture the information immediately. Sloppy or partial records at this point can create a cascade of problems for days if not decades to come. Even if the propagules that are collected never survive, the data (and the herbarium vouchers) will. The germplasm collector is responsible for collecting propagules, which typically are in the form of fruits, although cuttings and even entire seedlings are also an option. This person should have a good understanding of how to treat the collected material, from estimating seed viability and determining if the collection is warranted to selecting the right kind of bag. For example, oil-rich seeds like acorns can generate a lot of heat through respiration, so cloth bags are used to keep them from overheating. Rhododendron capsules, however, are best gathered in small glycine bags to prevent the tiny seeds from escaping. In addition to regular pruning shears, a set of pole pruners is very handy to have since they can extend an extra 10 to 12 feet (3 to 3.7 meters), often eliminating the need to climb trees. The herbarium voucher collector collects samples of plants that are then pressed and dried for future documentation and study. Collecting a voucher (often with several duplicates) is a critical part of the process because it captures the maternal plant's traits at reproductive maturity. Once deposited in a herbarium, the voucher will serve as evidence of what was growing in that location. During the trip, vouchers are kept in wooden plant presses, which, when tightly clamped down and ventilated, will yield high-quality pressed and dried specimens. Wooden presses can be used in the field but they are heavy and bulky, requiring extra time to pack and repack with each collection. Instead, some collectors use a plastic bag (the modern day version of a botanist's vasculum), which is a lot quicker to use. However, if not labeled properly specimens can get mixed up, and they can dehydrate by day's end. We prefer to use canvas field presses loaded with newsprint and corrugated cardboard. These presses are light weight, can be quickly loaded, and begin the pressing process immediately. By the end of the day, all of the specimens collected are transferred to the full-fledged wooden press, which can be tightened more effectively than the canvas press. While each of these activities may be assigned to a single point person, in reality it becomes a group effort. For instance, everyone might pitch in to call out associate species' names or other plant details to the trip recorder. Gather- MICHAEL DOSMANN A question often is asked: From how many mother plants do you need to collect? The answer depends. If the goal is to capture as much genetic variation of that species' population as possible, particularly if the seeds will be banked long-term in a repository, then the answer is \"many.\" The actual number depends on that species' breeding system (e.g., can it self-pollinate or is it forced to outcross), some technical assumptions based upon the population, including its size, and other details. The seeds from multiple mother plants can be bulked in the field (keeping track of how many were sampled), or each sample can be kept separate, which is important in cases such as the collection of conservation-status plants. If the goal is simply to produce a few living plants, then the focus may be less on capturing the full genetic variation of the population and just a few (or even just one) maternal plants can be sampled. Sometimes the sampling regime is dictated by realities in the field--there might just be one plant encountered, or maybe there is not enough time to hunt for and collect from a dozen separate individuals. And, for international collections, we ship seed back in smaller quantities (using \"small seed lot\" permits), so it may be prohibitive to collect from large numbers of maternal trees. Another question often posed is: How many collections are enough to call the expedition a success? This answer, too, depends on a number of factors. A garden that is in the expansion phase may have a lot of room to accommodate hundreds of new collections from a single expedition. However, for the Arnold Arboretum, where our goals for development are focused, fewer well-documented, high priority collections are more the norm. In fact, some trips may even be focused on just a single species. This can require practicing a bit of \"collection restraint\" to pass by plants that may be dripping with ripe fruits. However, it's also important to be open to \"opportunistic collections\" when a species of value--even though not a target species--is encountered. For example, while making our way towards a large The team made an \"opportunistic collection\" when this Acer griseum in southeastern Shaanxi in Septem- massive paperbark filbert (Corylus fargesii) was encountered in Shaanxi, China. ber 2015, our multi-garden team stumbled upon a massive Corylus fargesii. After genuflecting and hooting and hollering, we all agreed that it would be worth making a collection of this paperbark filbert for our respective institutions. Sometimes an opportunistic species is of value to just one of the gardens, prompting a discussion about whether the group should collect it or not. If the decision is made to collect it, even if just for one of the participants, it is important to treat it just like all of the other collections and properly catalog, document, and voucher it. Supplemental or extra collections made on the side can end up being a nightmare to track later on if not documented and assigned an official collection number! Also, sometimes an important species is found, but without any fruits. An official collection of it can still be made, since the herbarium voucher and documentation are of value, perhaps enabling a collector to return in the future. PHOTOS BY MICHAEL DOSMANN It's All in the Numbers Acquisition 9 KYLE PORT Larry Hufford prepares a voucher specimen of Douglas hawthorn (Crataegus douglasii) found east of Feather Creek in the St. Joe National Forest, Idaho. Herbarium specimens of nearly all of the 42 species collected during the 2015 North Idaho Expedition have been submitted to the Harvard University Herbaria for long term storage and scientific study. In Arkansas, during the 2014 Ozarks Expedition, Michael Dosmann (left) and Polly Hill Arboretum's Ian Jochams (right) arrange a voucher specimen of overcup oak (Quercus lyrata) in a canvas field press. ing fruits often falls to the full crew after other tasks are done, particularly if some of the seed extraction happens in the field. While typically all the participants are busy snapping photos, it is good to either assign the official task to one of the members, or at least make sure the recorder is jotting down which participant captured that collection's fullest complement of shots--habit, leaves, fruits, habitat, etc. A final photo swap at the end of the trip is a good way to cover all the bases. In some cases, X marks the spot and target species are found exactly where expected. But sometimes there's only the suggestion that the species occurs in the general area, requiring the entire group to pay close attention to the surrounding habitat for clues to the target's presence. For example, perhaps it is a droughttolerant tree more often found on the western or southern exposure of slopes, or it is a shadeloving shrub found in a mesic forest. Even with the team's expertise and a local collaborator in tow, plant identification in the field can sometimes be tenuous, so a trusted field guide is worth its weight in gold. These books can feel as heavy as gold when toted in a backpack, but luckily technology has come to the rescue. Michael reports that on multiple occasions he has been able to access the online version of the Flora of China from his phone, even while in remote sites in central China. What would E. H. Wilson think of that? And, since the team collects herbarium vouchers, further keying out can occur in the evenings or even after the expedition is concluded for some pesky and challenging species. EVENING ACTIVITIES At the end of the day's excursion all of the germplasm and herbarium vouchers are unpacked and double checked to make sure that collection numbers are properly assigned to each. Fruits are readied for drying or preparation, such as letting fleshy Viburnum fruits soak TIM BOLAND 10 Arnoldia 73\/4 KYLE PORT Acquisition 11 KYLE PORT KYLE PORT Propagules are bagged for shipment in specific ways: transplants, fleshy fruits, and cuttings are held in damp plastic bags while cones and seeds are placed in paper bags. Regardless of the container, handwritten notations including the name, collection number, and trip name are always applied and correspond to entries in a notebook or other form. These collections were made during the 2015 North Idaho Expedition. Dug from a nurse log beneath 400-year-old virgin timber in Hanna Flats, this Western red cedar (Thuja plicata) is the Arboretum's first wild collected accession from Bonner County, Idaho. KYLE PORT Hiking above 6,000 feet, our search for Larix lyallii was another incredible experience. In waning daylight, facing a two-mile hike out, we gathered cones from a particularly fecund specimen above lower Roman Nose Lake, Kaniksu National Forest, Idaho. After finishing the collection, we turned to take in the view and were treated to the aerial display of two golden eagles (the black specks in the upper center of this photo). 12 Arnoldia 73\/4 KYLE PORT Acquisition 13 part of the Animal and Plant Health Inspection Service (APHIS) of the United States Department of Agriculture (USDA). Once inspected by APHIS, they are then forwarded on to the Arboretum. Some collectors bemoan this step as unnecessary, and in some cases risky, since some items might be confiscated because of the presence of a pest or pathogen. For the Arboretum, this step actually provides peace of mind because it ensures that we are not importing the next invasive pest like Asian longhorned beetle or emerald ash borer. While the individual collections are documented by the recorder and ample photographs are taken, there is much that occurs during the trip that can only be captured in narrative form, a duty that falls to the designated trip journalist. Although such journaling requires extra time, it's valuable to have a trip participant capture the names of people met along the way, make broad observations of the different sites and regions visited, and chronicle the events that make plant exploration colorful such as meals, wildlife, and other cultural items. example, the ease of creating digital images in the field adds to post trip responsibilities. In times past, when glass plates and film were the only options (yes, we are that old), there were fewer images to annotate. Now it is not unusual to have hundreds if not thousands of images to label. In the end, documentation is a labor of love and we revel in its detailed complexity. Our passion for plants is manifest in the remarkable collections we steward. The Arnold Arboretum's 10-year Campaign for the Living Collections reaffirms a commitment to biodiversity discovery and the people engaged in plant exploration. In sharing our goals and experiences with you, we hope to inspire collaboration and exchange. If we range through the whole territory of nature, and endeavor to extract from each department the rich stores of knowledge and pleasure they respectively contain, we shall not find a more refined or purer source of amusement, or a more interesting and unfailing subject for recreation, than that which the observation and examination of the structure, affinities, and habits of plants and vegetables, affords. --Sir Joseph Paxton (1803 14 Arnoldia 73\/4 Acquisition 15 Clockwise from upper left: A striking stainless steel sculpture, Allium Spring Chorus by David Tonnesen, in McEuen Park, Coeur d'Alene. Once a hub for dairy operations, this barn now holds equipment at the University of Idaho Arboretum and Botanical Garden in Moscow. Specimens at Washington State University's Marion Ownbey Herbarium show the variation of leaf morphology in Acer glabrum var. douglasii, a species we collected. In St. Joe National Forest we collected seeds from snowbrush (Ceanothus velutinus), notable for its glossy foliage and fragrant white flowers. Master naturalist and Washington State University professor emeritus Steve Ullrich displays old man's beard (Usnea spp.), a lichenized fungi. 16 Arnoldia 73\/4 Acquisition 17 Clockwise from upper left: Michael Dosmann measures the diameter of an Acer griseum in Henan Province (photo by Kris Bachtell). Calcite deposits lead to the formation of amazingly colored iridescent pools, giving the Huanglong (Yellow Dragon) Valley in northern Sichuan its name. We made an opportunistic collection of this Hydrangea aspera (note the large pink sterile florets) during the 2015 NACPEC trip in Sichuan. Steep cliffs in Sichuan's Guangwushan Park were home to Acer griseum; although not visible in this image, one of the trees sampled grew just a few feet away from the precipice shown in the center-right of the image. Autumn color was seen on Fraxinus baroniana growing along the Jialing River in Gansu on the 2015 NACPEC trip. "},{"has_event_date":0,"type":"arnoldia","title":"BOOK EXCERPT: Saving the World's Deciduous Forests: Ecological Perspectives from East Asia, North America, and Europe","article_sequence":2,"start_page":18,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25602","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170af6b.jpg","volume":73,"issue_number":4,"year":2016,"series":null,"season":null,"authors":"Askins, Robert A.","article_content":"BOOK EXCERPT Saving the World's Deciduous Forests: Ecological Perspectives from East Asia, North America, and Europe Robert A. Askins Editor's Note: In this compelling book, biologist Robert A. Askins examines the history and ecology of Northern Hemisphere deciduous forest ecosystems in East Asia, North America, and Europe. These forests have a common ancient origin but have evolved in now widely separated regions for millions of years. Askins writes clearly on the similarities among and differences between the forests, including the threats to the plants and animals they contain and the challenge of developing effective conservation methods for these unique ecosystems. The excerpt presented here is from Chapter 5, \"Giant Trees and Forest Openings.\" Saving the World's Deciduous Forests: Ecological Perspectives from East Asia, North America, and Europe Robert A. Askins Yale University Press. 2014. 307 pages. ISBN: 978-0-300-16681-1 Book Excerpt 19 20 Arnoldia 73\/4 Book Excerpt 21 PHOTOS BY ROBERT A. ASKINS Clockwise from top: An imposing wooden gate stands at the entrance to the Strict Reserve in the heart of Poland's Bialowieza Forest. The understory seen here is quite open, with little between the tree canopy and vegetation on the forest floor. A view of the forest trail through the Strict Reserve. This photo shows the wide range of tree sizes (trunk diameters) in this old-growth forest where ancient trees stand along with middle-aged trees and young saplings. 22 Arnoldia 73\/4 Book Excerpt 23 24 Arnoldia 73\/4 Book Excerpt 25 a) 1.0 0.8 0.6 0.4 0.2 0.0 1.0 0.8 Betula 12 c) 10 8 6 4 12 8 6 4 2 0 Tilia Fraxinus Ulmus 10 8 6 4 2 0 12 8 6 4 2 0 1920 1930 1940 1950 1960 1970 1980 1990 2000 Populus Salix Picea 2 0 12 10 8 6 b) Picea abies 14 12 10 8 6 4 2 0 Tree recruitment (# *yr-1*ha-1) d) Carpinus 10 8 6 4 0.6 Quercus 0.4 0.2 0.0 20 15 10 5 0 1920 1930 1940 1950 1960 1970 1980 1990 2000 4 Acer Pinus 2 0 12 Ungulate density (ind. *km-2) 10 8 6 4 2 0 YEAR Alnus 2 0 e) Total Trees YEAR 26 Arnoldia 73\/4 ROBERT A. ASKINS Book Excerpt 27 28 Arnoldia 73\/4 Book Excerpt 29 30 Arnoldia 73\/4 Book Excerpt 31 32 Arnoldia 73\/4 ADAM WAJRAK Book Excerpt 33 (NOTES AND REFERENCES LISTED ARE FOR THIS EXCERPT ONLY) 34 Arnoldia 73\/4 Book Excerpt 35 "},{"has_event_date":0,"type":"arnoldia","title":"2015 Weather Summary","article_sequence":3,"start_page":36,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25601","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170af26.jpg","volume":73,"issue_number":4,"year":2016,"series":null,"season":null,"authors":"Pfeiffer, Sue A.","article_content":"2015 Weather Summary Sue A. Pfeiffer JANUARY began with seasonal temperatures. A warm front moved through on the 4th, bringing an inch of rain and a high temperature of 51 2015 Weather SUE PFEIFFER 37 No staff were lunching on the patio behind the Weld Hill Research Building on February 17. storm arrived on the morning of February 8th. The storm brought an additional 20 inches of snow, bringing the Arboretum's snow total to over 4 feet. Frigid conditions remained. With few places left to put the snow, the Arboretum crew spent many hours using large equipment to push snow banks back in order to clear roads and allow for access. Between the 1st and the 12th, 42 inches of snow fell, just surpassing the record for snowiest Boston February on record. The grounds were buried and many shrubs were completely covered, no longer visible under the thick quilt of snow. The final winter storm, a blizzard with high winds, dumped an additional 16 inches of snow on the 15th. By now, we were all exhausted from snow removal and were dreaming of spring. The idea of green grass, flowering snowdrops, and blooming magnolias seemed like an impossibility as snow banks of unprecedented size created a barrier between the roads and the collections. Sunny conditions returned and snow began to melt. There were a few smaller snowfalls over the following week and the temperature finally hit 41 Arnold Arboretum Weather Station Data 2015 Weather 39 week, peaking at 59 40 Arnoldia 73\/4 SUE PFEIFFER 2015 Weather 41 JULY was a dry month with seasonal temperatures. Intense thunderstorms on the 1st delivered three-quarters of an inch of rain in two separate downpours. Perfect summer conditions prevailed for most of the month; highs remained in the 70s and 80s, dipping to 69 42 Arnoldia 73\/4 KYLE PORT 2015 Weather ANDREW GAPINSKI 43 No jacket required: Arboretum Horticulturists Rachel Brinkman (left) and Sue Pfeiffer (right) spread soil amendments in the collections on the 10th of December, a month with temperatures that were well above average. NOVEMBER was a warm month and the 7th month of the year with below average precipitation. During the first week of November, temperatures hovered 15"},{"has_event_date":0,"type":"arnoldia","title":"Sensei: An Austrian Pine Forest Bonsai Comes to the Arboretum","article_sequence":4,"start_page":44,"end_page":44,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25603","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170b36f.jpg","volume":73,"issue_number":4,"year":2016,"series":null,"season":null,"authors":"Schneider, Stephen","article_content":"Sensei: An Austrian Pine Forest Bonsai Comes to the Arboretum Stephen Schneider ach dwarfed tree in the Arnold Arboretum's bonsai and penjing collection offers its own unique story. With the recent addition of ten new bonsai to this esteemed group, several interesting tales have been added to our archives. One of the new bonsai, an Austrian pine (Pinus nigra) forest, is of particular interest since it represents both a genus and a style that have never before been in our bonsai collection. Created and donated by Martin Klein of Andover, Massachusetts, this group of seven plants, meticulously sculpted over the past twenty-five years, takes its mound-like shape on what is known as a \"ciment fondu\" (a calcium aluminate cement) slab, also formed by the same artist. At first glance, one can't help but get the sense that this group of seven struggling stems represents nature on the edge of survival. Clearly competing for the most basic of resources-- light, water, nutrients--these tiny trees capture a snapshot of what their full-sized, earthbound counterparts contend with in the wild. This miniature forest receives daily care within the the Arboretum's comfortable urban setting, so it is all the more impressive that Martin's patience and steady hands created the successful illusion of ancient trees subjected to the vagaries of nature. Austrian pine, also known as black pine or European black pine, has a native range stretching from Austria to the Crimean Peninsula, south to Turkey and west to Morocco and Spain. It has been cultivated in the United States for at least 250 years. This two-needled pine grows 50 to 60 feet (15 to 18 meters) tall in the landscape and has characteristic dark green foliage and dark gray, furrowed bark. Austrian pine is quite adaptable, thriving under various soil and climate conditions (though unfortunately it is susceptible to diplodia tip blight, a fungal disease). It has a variety of uses including windbreaks, remediation plantings, and fastgrowing property screens. Its tolerance of dif- E ficult growing conditions makes it ideal for the pressures exerted through bonsai. The story of this bonsai forest began in 1991 when Martin purchased a bundle of ten Austrian pines from a local nursery for $25.00. Out of that bundle, seven plants were selected for the project. Emulating the work of bonsai master John Yoshio Naka, whom Martin had trained under during his early years of interest in bonsai, the concept of a forest began to take shape. Naka's famous bonsai, Goshin, a forest of eleven Foemina junipers (Juniperus chinensis `Foemina') on permanent display at the United States National Arboretum, inspired Martin to create a forest of his own using his newly acquired pines. The forest bonsai began in a training box where the young trees were pruned to develop trunk taper and wired for early form development. The forest then graduated to a Tokoname bonsai pot, and eventually moved to its final display on the ciment fondu slab. Mosses, lichens, and ferns were batted to the understory and have now all grown together, creating an emerald carpet speckled with many different textures and shades of green. At over four feet (1.2 meters) tall and with a spread of more than five feet (1.5 meters), this miniature forest creates a massive impact within the collection. On display for the first time this season, Arboretum visitors will find their imaginations challenged: To what distant land does this group of seven beckon them? Is this forest in a secluded mountain valley, or perhaps on a tiny island in the middle of a calm lake? As with all art, part of this bonsai's beauty lies in the perspective of the viewer. In memory of his former teacher John Yoshio Naka, Martin Klein has appropriately named this bonsai Sensei (teacher). As the fortunate recipient of this masterwork, the Arboretum will retain the name in the accession's passport data and proudly display our new treasure in the bonsai and penjing pavilion. Stephen Schneider is Director of Operations at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23448","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd1708528.jpg","title":"2016-73-4","volume":73,"issue_number":4,"year":2016,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Developing an Exemplary Collection: A Vision for the Next Century at the Arnold Arboretum of Harvard University","article_sequence":1,"start_page":2,"end_page":18,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25598","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170a76f.jpg","volume":73,"issue_number":3,"year":2016,"series":null,"season":null,"authors":"Pfister, Donald H.; Meyer, Paul W.; Hufford, Larry; Gapinski, Andrew; Donoghue, Michael J.; Boufford, David E.; Boland, Timothy M.; Dosmann, Michael S.; Friedman, William E.","article_content":"CAMPAIGN FOR THE LIVING COLLECTIONS Last autumn the Arnold Arboretum launched the Campaign for the Living Collections, an ambitious ten-year plan to expand the breadth of plant holdings and increase their scientific and horticultural value. Considerable thought and effort went into creating a document that guides the Campaign's mission. We present this important document here in its entirety for the benefit of Arboretum supporters, stakeholders, and colleagues. Additional articles covering aspects of developing our Living Collections will be featured in Arnoldia this year. Developing an Exemplary Collection: A Vision for the Next Century at the Arnold Arboretum of Harvard University William E. Friedman, Michael S. Dosmann, Timothy M. Boland, David E. Boufford, Michael J. Donoghue, Andrew Gapinski, Larry Hufford, Paul W. Meyer, and Donald H. Pfister T he Living Collections of the Arnold Arboretum of Harvard University not only support the Arboretum's mission by serving key research, education, and conservation roles, but in their entirety represent one of the very best examples of a historic Olmsted landscape. With some 15,000 accessioned plants, representing almost 4,000 unique taxa that include 2,100 species, the Living Collections of the Arnold Arboretum remain a major destination for those who study and enjoy woody plants. Of the accessions brought to the Arboretum from elsewhere, 44% are of wild origin, hailing from over 60 different temperate countries. Another 39% are of cultivated origin, including pedigreed hybrids, nursery-origin introductions, and accessions from other gardens. This historic interplay between taxonomic, floristic, and cultivated diversities has resulted in one of the most comprehensive and heavily documented collections of temperate woody plants in the world. The living collections are central to the Arnold Arboretum--all research, education, and conservation initiatives are driven by them. And yet, without strategic planning for collections development, these collections are at risk of losing their prominence. In advance of the Arboretum's sesquicentennial in 2022, it is time to put forward a set of initiatives to simultaneously preserve its singular legacy and secure its future. This plan, to be enacted over the next decade, will thus serve to shape and define the Living Collections of the Arnold Arboretum for the coming century. Values and Aspirations Throughout the Arboretum's history, the vitality and strength of the collections--and the institution as a whole-- can be attributed to an adherence to four essential tenets. The Arboretum's collection of Malus (apples and crabapples) currently holds 426 individual plants from 310 accessions comprising 159 taxa, many of which grow on Peters Hill, seen here. It has remained a traditional arboretum, with the Living Collections continually and almost exclusively focused on temperate woody plants. It has been committed to collections-based woody plant scholarship, particularly in recent years with the significant expansion of on-site research associated with construction of the Weld Hill Research Building. The Arboretum landscape remains true to the vision of Frederick Law Olmsted's design, through keen awareness of its role as a public garden and landscape. Lastly, the Arnold Arboretum has long invested in active curation and collections management, which in turn has fostered and enabled its research and preservation enterprises. Coupled to these principles are aspirations that the Living Collections Advisory Board (Tim Boland, David Boufford, Michael Donoghue, Larry Hufford, Paul Meyer, and Don Pfister), in collaboration with William (Ned) Friedman (Director), Michael Dosmann (Curator of Living Collections), and Andrew Gapinski (Manager of Horticulture), advanced during its 2013, 2014, and 2015 meetings. These recommendations to ambitiously strengthen the collections of the Arnold Arboretum articulate a set of guidelines that define targets for active collections development. In addition, these recommendations clearly circumscribe overarching principles that will help ensure that the Arnold Arboretum remains at the leading edge of botanical garden collections development. Enacting the Agenda and Creating a List of Desiderata Four principles will shape a unique identity for the Arboretum's ongoing and future roles in collections-based research, teaching, and public horticulture: (1) scholarship associated with comparative biology, from genomics to environmental change; (2) ex situ conservation and study of endangered temperate woody plant taxa; (3) strengthened species representation within key priority genera; and (4) successful cultivation of taxa currently or historically perceived as marginally hardy in Boston. Shared among these is the primary importance of a university-based organismic collection and public garden with the potential to uniquely reveal the complexities of nature. This deeply held notion expresses the intrinsic value that an individual accession or a suite of accessions possesses that enables a scholar to unravel (and share with the world) a taxon's mysteries. This extends to naturally occurring taxa as well as unique cultigens, honoring the Arboretum's long history of cultivating and comparing both, side-by-side. This core philosophy underpins all others. In support of these priorities for collections development, six goals have been used to specifically create and prioritize a list of desiderata for acquisition and accessioning into the organismic collections of the Arnold Arboretum over the next decade (see page 15). These six goals NANCY ROSE 4 Arnoldia 73\/3 NANCY ROSE le m en Le nt siae ag o* Pu nc ta Eu ta vi bu Ps rnu m eu * do tin So us le * no tin V. us lu * te To scen m en s V. tosa am * p Ur li c at ce um ol at Ti a nu s Viburnum Plant Phylogeny When explaining the relatedness of organisms, scientists often construct a phylogeny, or \"family tree,\" that portrays the organisms' evolutionary history. This figure depicts taxa (such as species, genera, families, etc.) on the tips or ends of the branches. Each taxon placed at the end of a branch is paired with its closest relative; the two can be traced back to a shared node that represents their common ancestor. That branch, in turn, can be traced back further to another ancestor that is common to it and another \"sister\" branch, ad infinitum. A branch that includes the common ancestor and all of its descendants is referred to as a clade. In contemporary phylogenetic systematics, a plant family's phylogeny should comprise individual clades of species that form distinct genera. Genera themselves comprise clades of species that share a common ancestor, and may receive special names. Organizing related species into clades is useful when studying large genera and families, particularly if the group is supported by a well-developed and accepted phylogeny. In such cases scientists look at all of the clades that occur across the breadth of the phylogeny but only need to sample one or two species per clade. We have taken the same approach to collections development at the Arboretum, where comprehensive or total species composition within a genus is not a goal but phylogenetic breadth (i.e., representation of as many clades as possible) is. The genus Viburnum (comprising some 165 species) is one such example. Decades of work in Michael Donoghue's laboratory has yielded a well-supported Viburnum phylogeny containing 19 terminal clades of closely related species (above, modified from Clement et al. 2014). Ten of these clades (shown in blue) are already represented in the Arboretum's collections by at least one species; acquisition of a few more species (* ) improves their representation. Although the remaining clades contain species not suited to New England (most are tropical), four clades (shown in red) contain a few marginally-hardy species that are worth trialing. Adding even just one species from a clade (e.g., V. davidii in the Tinus Clade) adds a completely new evolutionary lineage to the collections. Clement, W. L., M. Arakaki, P. W. Sweeney, E. J. Edwards, and M. J. Donoghue. 2014. A chloroplast tree for Viburnum (Adoxaceae) and its implications for phylogenetic classification and character evolution. American Journal of Botany 101: 1029 6 Arnoldia 73\/3 Genome Sequencing Brings New Knowledge Genome sequencing, which allows the entire string of DNA code to be read and interpreted for almost any species on earth, has the potential to link specific biological characteristics to specific genes and their genetic variants. Fifteen years ago, it cost nearly $100 million to sequence a genome and today, this cost is rapidly approaching $1,000. Thus, in the last decade, woody plant species as diverse as grape, poplar, pine, willow, oak, and birch have had their genomes sequenced, so that their biological traits can be related directly to their genes. Many more species of trees, shrubs, and lianas are in the pipeline for genomic sequencing. In the spring of 2015, the Arnold Arboretum hosted an international symposium on the genomics of forests and trees that brought together 80 of the leading scholars in this emerging world of DNA information. One critical challenge that came up repeatedly in discussions with participants is that those organisms whose genomes have been or are going to be sequenced are at high risk of being lost since there is no formal plan for perpetual stewardship. Incredibly, there is no home in the entire world for the unique individual plants whose genomes have been sequenced. The Arnold Arboretum has a perfect combination of world-class horticulture and world-class laboratory facilities to pioneer a \"genomic type collection\" of temperate woody plants, and will care in perpetuity for these incredibly special individual plants whose genomes have been sequenced. This genomic type reference collection will help tell us how trees will respond to climate change, how water moves from the soil to the top of the crown, how plants respond to biotic (herbivores) and abiotic (climate extremes) stresses. As research comes to rely more heavily on sequenced genes to advance discovery, the ability to reference the living counterpart of this DNA--either through the plant itself or its direct lineage--will make the Arnold Arboretum an essential resource for this work. raphy, ecophysiology, and phylogenetics, acquiring disjunct species associated with clades of eastern North American woody plants will be a priority. Although the majority of these disjunct taxa are of Asian origin, disjunct taxa from Europe and the Southern Hemisphere (although fewer in number) will also be sought. 6. Reassert horticultural leadership in accessioning marginally hardy species As established at its founding in 1872, the Arboretum will cultivate \"all the trees, shrubs, and herbaceous plants, either indigenous or exotic, which can be raised in the open air.\" In the spirit of exploration and experimentation, the Arboretum has continually acquired germplasm of marginally hardy taxa to be coaxed into cultivation, despite and against all odds. To ensure that the Arboretum stays at the cutting edge of plant introduction (especially in a world of rapid environmental change), it must seek out, acquire, and test untried species for growth on the grounds. Importantly, identification of new \"marginal\" taxa should be coupled with targeted field collections of germplasm from parts of the taxon's natural range that are likely to predispose such accessions to ultimate success on the grounds. In light of predicted climate change scenarios over the next century in Massachusetts, the marginally hardy taxa of today are likely to be well positioned to grow and thrive at the Arnold Arboretum in the future. With all of this in mind, a list of desiderata comprising roughly 400 separate taxa has been developed, where each target for acquisition falls into one and often multiple prioritized categories or themes based on the guiding prin- 8 Arnoldia 73\/3 ANTHONY S. AIELLO Developing an Exemplary Collection 9 Table 1. High-priority targets for future acquisition comprise 395 separate taxa that represent 385 species among 145 distinct genera (see List of Desiderata). Each distinct target is linked to one or more of the priority goals put forward by the Collections Advisory Board. This table summarizes those core themes and other basic statistics. New species for the collections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177 marginally hardy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 New genera for the collections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 marginally hardy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Taxa already represented in the collection by living lineages . . . . . . . . . . . 218 ... but not of wild provenance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 ... of wild provenance but requiring additional wild lineages z . . . . . . 121 Principal categories y: Conservation concern x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 PCN genera w . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Robust genera v . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Disjunct genera u . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Acquisition targets' regions of origin: Eastern Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225 Eurasia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133 South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 z Includes taxa where numerous wild-origin lineages are required, often for conservation purposes, or to repatriate known Arboretum lineages from other repositories and gardens. y Categories are not mutually exclusive; many target species occur in multiple categories. x Those taxa to which a conservation ranking has been given. w Plant Collections Network: Acer, Carya, Fagus, Stewartia, Syringa, and Tsuga v Carpinus, Forsythia, Ginkgo, and Ostrya u Only pertains to species occurring in Cornus, Diervilla\/Weigela, Hamamelis, Hydrangea, Magnolia, Taxus, and Viburnum 10 Arnoldia 73\/3 Developing an Exemplary Collection 11 WILLIAM E. FRIEDMAN WILLIAM E. FRIEDMAN The Carya collection is one of six Plant Collections Network collections currently at the Arboretum. Seen here, fruits of shagbark hickory (C. ovata 22868-N) and bud scales and new shoots of shellbark hickory (C. laciniosa 806-87-C). nial in 2072, the Arboretum community will be able to look back and pinpoint this campaign as one that guaranteed and secured a future where the Arnold Arboretum became the single most important living collection for those who study and enjoy temperate woody plants. Developing an Exemplary Collection-- Metrics and a List of Desiderata Over the past decade, the total number of deaccessions has significantly exceeded the total number of new accessions planted at the Arnold Arboretum. On average each year, roughly 250 new accessions are successfully added to the collections. In order to achieve the goals described in this plan to secure the long term preeminence of the Living Collections of the Arnold Arboretum, acquisition and successful propagation of new and diverse germplasm will dominate the early phases of this campaign. The second phase of this ambitious program will then leverage the accomplishments of plant production efforts to move new accessions onto the grounds. We anticipate that at the peak of this second phase close to 300 new accessions will be added to the permanent Living Collections annually. The new leadership in collections-based horticulture (Andrew Gapinski) and in plant production (Tiffany Enzenbacher), along with the recent significant improvement in the management and job circumscriptions of horticulturists charged with the day-to-day care of the living collections, will ensure that this plan is successful. The horticultural team will be charged, under the leadership of the Curator of Living Collections (Michael Dosmann) and Director (William Friedman), with the creation of a specific set of plans and targets to fully enact the vision of this decadal campaign for the next century of the Arnold Arboretum's history. List of Desiderata: Priority Genera (Plant Collections Network, Robust, and Disjunct Genera) Highlighted are 17 genera that will receive focused investment in review and future acquisition. In fact, over 40% of the listed taxa are within this group. Although the collection goals for each of these are similar, there are some subtle differences in the levels at which each target should be represented. 12 Arnoldia 73\/3 Developing an Exemplary Collection 13 Living Collections are to be considered fully synoptic and representative of Earth's temperate woody flora, over 175 species and 40 genera not represented will be targeted and acquired. Acquisition of the rare is essential, and the list of desiderata includes over 50 MICHAEL S. DOSMANN targets with specific conservation value as measured by a NatureServe ranking of G1 to G3 (critically imperiled, imperiled, vulnerable) or its general equivalent. Documentation is the most reliable predictor of accession value, and many targets were selected to fill documentation gaps. One-fourth of the target taxa on the list are already growing in the Living Collections, yet are represented by accessions that are not of wild origin. It is essential that these \"finally wild\" targets are integrated into the Living Collections to displace specimens that have poorer (or no) documentation. Another third of the target taxa already exist in the collections and include at least one wild-origin accession. However, additional wild lineages are required to broaden the diversity of that taxon's lineage pool. An additional one-fourth of the target taxa either have not been tried at the Arboretum in the past, or they were planted but failed to establish because they were not sited in a suitable microclimate or for other reasons. In the spirit of exploration, hardiness testing, and plant introduction, the Arboretum will trial these taxa. MICHAEL S. DOSMANN Genomic Type Specimens Initially, the number of accessions associated with published genomic data will likely number in the single digits. However, with time, improvements in DNA sequencing technology and data analysis will likely yield dozens, if not hundreds of temperate woody plant taxa with sequenced genomes. The Living Collections Advisory Board will be charged with assessing which taxa should be added to the genomic type specimens of the Arboretum. Initially, attention to phylogenetic breadth Ostryopsis davidiana (top) and Acer caudatum ssp. multiserratum (bottom) are among the species on the list of desiderata. The specimens seen here were photographed in China during the 2010 NACPEC expedition. 14 Arnoldia 73\/3 List of Desiderata Family Actinidiaceae Adoxaceae Taxon Actinidia chinensis Viburnum atrocyaneum Viburnum bracteatum Viburnum buddleifolium Viburnum carlesii Viburnum cylindricum Viburnum davidii Viburnum ellipticum Viburnum farreri Viburnum foetidum Viburnum ichangense Viburnum kansuense Viburnum lantanoides Viburnum microcarpum Viburnum mongolicum Viburnum obovatum Viburnum plicatum ssp. tomentosum Viburnum sieboldii Viburnum tinus Viburnum urceolatum Viburnum wilsonii Liquidambar styraciflua Cotinus szechuanensis Asclepias incarnata Trachelospermum difforme Aralia nudicaulis Araucaria araucana Aristolochia tomentosa Nandina domestica Alnus glutinosa Alnus maritima Betula alleghaniensis Betula alnoides Betula chichibuensis Betula humilis Betula lenta Betula luminifera Betula maximowicziana Betula murrayana Betula pumila Betula schugnanica Betula uber Carpinus caroliniana Carpinus cordata Carpinus coreana Carpinus henryana Carpinus polyneura Ostrya japonica Ostryopsis davidiana Cupressaceae Cyrillaceae Daphniphyllaceae Elaeagnaceae Ephedraceae Family Bignoniaceae Buxaceae Taxon Campsis grandiflora Pachysandra procumbens Pachysandra terminalis Sarcococca hookeriana var. digyna Opuntia humifusa Aphananthe aspera Celtis tenuifolia Kolkwitzia amabilis Leycesteria formosa Weigela coraeensis Weigela maximowiczii Weigela middendorffiana Euonymus fortunei Euonymus obovatus Cercidiphyllum magnificum Sarcandra glabra Alangium platanifolium Aucuba japonica Cornus alternifolia Cornus canadensis Cornus florida Cornus foemina Cornus nuttallii Cornus quinquinervis Cornus rugosa Cupressus nootkatensis Cliftonia monophylla Daphniphyllum macropodum Elaeagnus multiflora Ephedra distachya Ephedra equisetina Ephedra gerardiana Ephedra intermedia Ephedra likiangensis Ephedra major Ephedra minuta Ephedra monosperma Ephedra przewalskii Ephedra rituensis Ephedra sinica Ephedra viridis Agarista populifolia Eubotryoides grayana Gaultheria miqueliana Gaultheria procumbens Kalmia cuneata Cactaceae Cannabaceae Caprifoliaceae Celastraceae Cercidiphyllaceae Cloranthaceae Cornaceae Altingiaceae Anacardiaceae Apocynaceae Araliaceae Araucariaceae Aristolochiaceae Berberidaceae Betulaceae Ericaceae 16 Arnoldia 73\/3 Developing an Exemplary Collection 17 Family Oleaceae Taxon Chionanthus virginicus Forsythia europaea Forsythia japonica Forsythia mandschurica Forsythia suspensa Forsythia togashii Fraxinus americana Fraxinus nigra Fraxinus pennsylvanica Fraxinus quadrangulata Osmanthus americanus Syringa afganica Syringa emodi Syringa josikaea Syringa julianae Syringa komarowii Syringa komarowii ssp. reflexa Syringa mairei Syringa meyeri Syringa oblata Syringa oblata ssp. dilatata Syringa pinetorum Syringa pinnatifolia Syringa protolaciniata Syringa pubescens ssp. microphylla Syringa reticulata Syringa sweginzowii Syringa tibetica Syringa villosa Syringa vulgaris Syringa wardii Syringa yunnanensis Meconopsis integrifolia Abies alba Abies koreana Larix laricina Larix lyalii Larix occidentalis Larix potaninii Picea abies Picea omorika Pinus albicaulis Pinus cembra Pinus lambertiana Pinus monophylla Pinus virginiana Pseudolarix amabilis Tsuga canadensis Tsuga caroliniana Tsuga chinensis Tsuga diversifolia Tsuga dumosa Tsuga forestii Family Taxon Tsuga formosana Tsuga heterophylla Tsuga mertensiana Tsuga sieboldii Platanaceae Ranunculaceae Platanus orientalis Clematis akebioides Clematis tangutica Clematis viridis Ziziphus jujuba Amelanchier nantucketensis Crataegus distincta Crataegus harbisonii Crataegus latebrosa Crataegus perjucunda Crataegus phaenopyrum Cydonia oblonga Exochorda korolkowii Exochorda racemosa Exochorda serratifolia Malus angustifolia Malus coronaria Malus florentina Malus ioensis Malus komarovii Malus prunifolia Malus transitoria Mespilus canescens Neviusia albamensis Physocarpus opulifolius Prunus apetala Prunus cyclamina Prunus maritima Prunus sargentii Rosa chinensis Rosa hugonis Rubus paludivagus Rubus prosper Rubus saltuensis Sorbaria kirilowii Sorbus domestica Sorbus wilsoniana Spiraea sargentiana Adina rubella Emmenopterys henryi Mitchella undulata Populus grandidentata Populus tremuloides Salix jejuna Salix turnorii Buckleya angulosa Buckleya distichophylla Rhamnaceae Rosaceae Papaveraceae Pinaceae Rubiaceae Salicaceae Santalaceae 18 Arnoldia 73\/3 "},{"has_event_date":0,"type":"arnoldia","title":"Legacy Trees of Ernest Henry Wilson and John George Jack in Nikko, Japan","article_sequence":2,"start_page":19,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25599","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170ab28.jpg","volume":73,"issue_number":3,"year":2016,"series":null,"season":null,"authors":"Ohkubo, Tatsuhiro; Aizawa, Mineaki","article_content":"Legacy Trees of Ernest Henry Wilson and John George Jack in Nikko, Japan Mineaki Aizawa and Tatsuhiro Ohkubo ikko, in Tochigi Prefecture, about 80 miles (130 kilometers) north of Tokyo, has been among the most famous resorts in Japan for foreign visitors since the Meiji era (1868 20 Arnoldia 73\/3 Legacy Trees 21 available online in the digital archives of the Arnold Arboretum (directions for searching the database can be found at: http:\/\/www. arboretum.harvard.edu\/library\/image-collection\/botanical-andcultural-images-of-eastern-asia\/search-the-image-database\/). E. H. Wilson visited Japan in 1914 and traveled throughout the country. According to his itinerary, he visited Nikko and photographed its trees and landscapes in May and October of that year. In 1905, nearly a decade before Wilson's visit, J. G. Jack also stayed in Nikko. He was interested in natural forests and forestry in Japan, and his photographs thus give us a glimpse into the forestry and lumber industries of Japan at that time. From 2007 to 2015, we traced the footprints of these visitors, and in some places we encountered the trees as Wilson and Jack had seen them in the past; the trees have kept their living witness for over a hundred years. We present here comparison photographs of these trees from the past and present. Japanese larch and Japanese arborvitae near Yumoto In the photographs that Wilson made near Yumoto on October 16, 1914, we looked at the probable view from Lake Karikomi, which takes an hour to reach by walking from Yumoto, and concluded that he must have traveled along a footpath that had been drawn from Yumoto to the lake in an old Japanese topographical map, published in 1915. While walking along the footpath, we immediately encountered an inspirational sight. The massive trunk of a Japanese larch (Larix kaempferi) that we had previously seen was standing in plain view. This tree was the one from Wilson's 1914 photograph, which appeared as plate XVI in his book The Conifers and Taxads of Japan, published in 1916. Information from the label on the back of the photo mount indicated that the tree's trunk was 12 feet (366 centimeters) in girth when originally photographed. In 2008, the tree was 12.5 feet (382 centimeters) in girth and 95 feet (29 meters) in height, demonstrating slight growth. Nikko is famous for its native larch, and this tree is a fine representative of the larch in this region. In Wilson's larch photo, a tree with multiple trunks can be seen just behind the man posing next to the larch. This tree is a Japanese arborvitae (Thuja standishii), photos of which also appear in his book (plates LII and LIII). It is surprising that a branch that inclined left and downward, as well as the spatial form of the tree behind the Japanese arborvitae, were in precisely the same positions as they were a hundred years ago (AEE-03655 and view in 2008). Now the tree is 15.9 feet (484 centimeters) in girth and 76 feet (23 meters) in height, while in 1914 it was 11 feet (335 centimeters) in girth and 45 feet (14 meters) in height. After an hour of walking, we reached the lakeside of Lake Karikomi. We tried to locate a massive Japanese bird cherry (Prunus ssiori), with a girth of 9 feet (274 cm) in 1914, that had stood at the water's edge. Although we were able to track down the view from behind the tree, the tree is no longer in existence. 22 Arnoldia 73\/3 Legacy Trees 23 Japanese yews and another Japanese arborvitae around Lake Chuzenji Next, we examined the trees around Lake Chuzenji, which is located at the foot of Mount Nantai, a volcano that rises to a height of 8,156 feet (2,486 meters). Because the embassy villas of several countries were located around the lake during the Meiji era, foreign visitors may be familiar with this area. The Nikko Futarasan-Jinja Chugushi Shrine is located on the north side of Lake Chuzenji. Jack and Wilson both visited this shrine. Jack photographed two tall trees (AEE-00175 and 00176) in this shrine. The two tall trees are now lost. According to the information from the label of Jack's photo, these trees were Japanese umbrella pine (Sciadopitys verticillata). However, we wonder if these trees may have been Japanese soft pine, Pinus parviflora var. pentaphylla, considering their shapes and the non-native status of Sciadopitys verticillata in Tochigi. There is the possibility, though, that Sciadopitys verticillata was purposely planted at the shrine, as is occasionally observed in east and west Japan. In these photos (AEE-00175 and 00176), gravel sediment can be seen on the ground, which represents the remains of a massive landslide on Mount Nantai caused by a typhoon in September 1902. The shrine buildings and two tall trees in front of the building barely escaped the damage. In the photo AEE-00175, a Japanese yew (Taxus cuspidata) can be seen located to the right of the two tall trees. This tree is still alive and it is now designated as a natural monument of Tochigi Prefecture. The age of the tree, as estimated from the age of a broken large branch, is more than 1,000 years. The tree was last measured at 12.8 feet (390 centimeters) in girth and 62.3 feet (19 meters) in height in a 1988 investigation by the Ministry of Environment of Japan. Another massive Japanese yew photographed by Jack (AEE00180) is still alive behind a main building of the shrine. Jack indicated that this tree was more than 3.5 feet (107 centimeters) in diameter and 50 feet (15 meters) in height in 1905. In 2015, the tree was 3.9 feet (119 centimeters) in diameter and 68.9 feet (21 meters) in height. When we observed the tree in 2010, its trunk was covered with a plastic net for protection against bark stripping by sika deer; in Nikko and throughout Japan, bark stripping by sika deer is now among the most serious problems facing forest ecosystem and forestry. Japanese yew is one of the favorite plants of sika deer. The Tachiki-Kannon temple is located on the east side of Lake Chuzenji. A wooden standing statue of the eleven-faced, onethousand-armed Kannon Bosatsu is enshrined in the main hall of the temple. This Kannon Bosatsu was carved into a standing tree with roots (\"Tachiki\" in Japanese), which is why the temple is called \"Tachiki-Kannon.\" This Kannon was said to be carved in 24 Arnoldia 73\/3 Legacy Trees 25 M. AIZAWA M. AIZAWA The shrine buildings, with a sacred Taxus cuspidata surrounded by a red wooden fence (seen lower right), at the Nikko Futarasan-Jinja Chugushi Shrine, Lake Chuzenji, on July 10, 2010. A front view of the sacred Taxus cuspidata surrounded and protected by a red wooden fence at the Nikko Futarasan-Jinja Chugushi Shrine, Lake Chuzenji, on July 10, 2010. M. AIZAWA Taxus cuspidata, Lake Chuzenji, on August 10, 1905, photographed by J. G. Jack (AEE00180 from the Arnold Arboretum Archives). Probably the same Taxus cuspidata at the Nikko Futarasan-Jinja Chugushi Shrine, by Lake Chuzenji on November 4, 2015. Note the plastic mesh fencing to deter sika deer from stripping bark from the trunk. 26 Arnoldia 73\/3 Legacy Trees 27 M. AIZAWA \"Chamaecyparis obtusa,\" near \"Kwannon Temple,\" Lake Chuzenji, on May 29, 1914, photographed by E. H. Wilson (AEE-03424 from the Arnold Arboretum Archives). The tree is actually Japanese arborvitae (Thuja standishii). Thuja standishii (not Chamaecyparis obtusa) at the Tachiki-Kannon temple, Lake Chuzenji on July 10, 2010. the late eighth century. The temple was originally located next to the Nikko Futarasan-Jinja Chugushi Shrine on the north side of Lake Chuzenji. However, the temple was destroyed by the debris flow from the above-mentioned landslide in 1902 and the Kannon was swept away into the lake. The statue was subsequently beached, and the temple was reconstructed in its beached location in 1913. The tree that stood out most conspicuously at the temple was another Japanese arborvitae that Wilson photographed on May 29, 1914 (AEE-03424, 03425). Wilson described this tree as a Hinoki cypress (Chamaecyparis obtusa), but it is actually a Japanese arborvitae. In 2010, the tree was 19.8 feet (603 centimeters) in girth and 103 feet (32 meters) in height, indicating slight growth from the 19 feet (579 centimeters) in girth and 90 feet (27 meters) in height observed in 1914. According to the giant tree database of Japan, only a few Japanese arborvitaes in the country have girths greater than 32.8 feet (10 meters). The biggest specimen of the species in Tochigi Prefecture is reported to be 21.0 feet (640 centimeters) in girth, located in Yunishigawa, Nikko. The Japanese arborvitae that Wilson photographed must now be comparable to that recorded tree. 28 Arnoldia 73\/3 Legacy Trees 29 Japanese cedar (Cryptomeria japonica), Nikko, on May 19, 1914, photographed by E. H. Wilson (AEE-03754 from the Arnold Arboretum Archives). Photograph from the same position as that of the left photograph on January 13, 2007. Forests after the fires of the 1890s near Yumoto Finally, we introduce our recent study (Ishida et al. 2013), which provided evidence for forest fires near Yumoto in the Meiji era and was inspired by Jack's photographs. Jack photographed in Yumoto two views of forests after fires, entitled \"Japan-Forestry\" (AEE-00202 and 00203). According to the information from the labels, forest fires after lumber harvest had occurred on the south slopes, which were originally covered with fir (Abies), hemlock (Tsuga), larch (Larix), birch (Betula), oak (Quercus), and other trees, and had burned from May 15 to June 15 around 1893. Locating these burned forests bolsters the understanding of the forest dynamics for the century after a fire. Using Jack's photographs, we tried to locate the sites when the deciduous species were leafless, giving us a better view of the skyline and geological features, but we did not identify the locations in a year of searching. But then, when we looked at an old Japanese topographical map drawn in 1915, we noticed that it indicated the presence T. OHKUBO 30 Arnoldia 73\/3 Legacy Trees 31 ISHIDA ET AL. 2013 A (AEE-00202) and D (AEE-00203), views of previously burned forest on slopes in Yumoto photographed on August 11, 1905, by J. G. Jack; B and E, birds-eye views of the slopes constructed using Kashmir 3-D software; C and F, current views of the slopes with maturing forests on December 1 and December 4, 2010. Acknowledgements We are grateful to the people who helped us locate these trees in Nikko, and we also thank R. Primack for encouraging us to locate the Japanese trees of the Arnold Arboretum archives. The works of Primack and Ohkubo (2008) and Flanagan and Kirkham (2009) inspired us to write this article. References Flanagan, M. and T. Kirkham. 2009. Wilson's China: A century on. Richmond, Surrey: Kew Publishing, Royal Botanic Gardens, Kew. Ishida, Y., M. Aizawa, and T. Ohkubo. 2013. Evidence of forest fires in the Meiji era in the mountainous region of Nikko on the basis of charcoal particle analysis. Japanese Journal of Forest Environment 55: 1"},{"has_event_date":0,"type":"arnoldia","title":"Woodland Restoration: 30 Years Later","article_sequence":3,"start_page":32,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25600","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170ab6d.jpg","volume":73,"issue_number":3,"year":2016,"series":null,"season":null,"authors":"McCoy, Emily; Mandel, Lauren","article_content":"Woodland Restoration: 30 Years Later Lauren Mandel and Emily McCoy W alking along the wooded trails in Loantaka Brook Reservation, Morristown, New Jersey, it's hard to believe that a 36-inch-diameter natural gas pipeline lies underfoot. The mature oak-beech forest overhead and native herbaceous understory don't suggest that this was a construction zone, nor do the runners or cyclists zipping by appear to notice. The Algonquin Gas Transmission pipeline, which transports 2.74 billion cubic feet of natural gas per day between New Jersey, New York, and parts of New England (Spectra Energy 2015), runs directly through the reservation. Its installation through the wooded landscape began 30 years ago, in 1986. Continual site monitoring suggests that the use of minimally invasive construction methods during the pipeline's installation, paired with innovative habitat restoration techniques and stewardship guidelines, have resulted in the long-term ecological success of this sensitive, post-construction landscape. THE SITE The reservation, which is used primarily for walking, running, cycling, horseback riding, and cross-country skiing, lies 25 miles due west of Manhattan, within the northernmost portion of New Jersey's Great Swamp watershed. The ALL IMAGES BY ANDROPOGON ASSOCIATES UNLESS OTHERWISE NOTED Loantaka Brook Reservation, Morristown, New Jersey, contains diverse plant communities including several forest and meadow types. To limit ecological disruption, a natural gas pipeline installed through the reservation in 1986 was carefully routed and constructed. Here, the pipeline alignment is being marked. Woodland Restoration 33 As part of the preparation for the pipeline installation in 1986, a plant community inventory that mapped the various meadow, woodland, and forest species assemblages was completed. (Color coding added for this article.) Loantaka Brook, the reservation's namesake, flows from just north of the reservation down into the neighboring Great Swamp National Wildlife Refuge, where it joins other flows that discharge into the Passaic River. The reservation falls within the Glaciated Reading Prong\/ Hudson Highlands regional ecosystem, with local soils containing glacial lake bottom deposits of fine sand and silt with clay. Traversing this region, the 1,129-mile-long Algonquin pipeline connects to the mammoth Texas Eastern Transmission pipeline, which together provide approximately one-third of the continental United States with natural gas transportation infrastructure. The pipeline route was originally slated to run near a residential neighborhood in Morristown, but instead the Federal Energy Regulatory Commission mandated that the infrastructure be rerouted through the 744-acre reservation. The reservation contains diverse plant communities that respond to the site's topography and land use history. In the 1980s, young forest dominated the reservation, with sizeable lowland woodland areas, primarily consisting of red maple (Acer rubrum), oak (Quercus spp.), and ash (Fraxinus spp.), all of a similar age. Upland woodlands were also present, with dominant oak, hickory (Carya spp.), black birch (Betula lenta), tulip poplar (Liriodendron tulipifera), and red maple species. Mature forest stands within the reservation consisted of mixedage, stable plant associations with a stratified forest structure. Upland forest communities contained large populations of American beech (Fagus grandifolia), tulip poplar, and red oak (Q. rubra), while lowland forests were dominated by red maple, white oak (Q. alba), and ash. The site also contained meadows that, in the 1980s, were in an early successional state that exhibited herbaceous perennials, pioneer woody shrubs, and young trees. Upland meadow areas were dominated by forbs, grasses, multiflora rose, goldenrod, and ferns, while wet meadow 34 Arnoldia 73\/3 Woodland Restoration 35 This figure shows details of habitat protection measures instituted during the 1986 pipeline installation. compressing the soil's air-filled pores (Sch 36 Arnoldia 73\/3 Woodland Restoration 37 One year after pipeline installation, the forest construction zone looked remarkably undisturbed. 35-foot-wide construction zone, while invasive species were eliminated. Approximately 25 feet of wooded area beyond the construction zone, in both directions, was considered a Park Commission zone of special management. MONITORING METHODS Various types of landscape monitoring occurred immediately before the pipeline's 1986 construction, periodically from the late 1980s through the late 2000s, and then with more rigor in 2013. Landscape architect Leslie Sauer recalls that in the 1980s landscape performance monitoring was not common practice and, because of this \"analysis paralysis,\" the pipeline owner did not agree to allocate funds for a formal investigatory effort (Sauer 2015). Informal monitoring was therefore deployed initially to inform the designers of the effectiveness of the new design, construction, and stewardship strategies. The site investigations performed in 1986, in preparation of the pipeline installation, included a plant community inventory that mapped the various meadow, woodland, and forest species assemblages. Additionally, a tree inventory and shrub valuation within and adjacent to the realigned pipeline corridor helped designate each woody plant for protection or removal. One year after construction, the landscape architect visually monitored the site. For nearly three decades to follow, monitoring methods consisted primarily of observation and photographic analysis during periodic site visits. In order to assess the ecological success of the implementation strategies after nearly 30 years, the integrative research department at Andropogon Associates performed vegetation and soil sampling in June 2013. The assessments aimed at understanding how the pipeline construction methods affected the biodiversity and soil health of the forest within the construction zone in comparison to areas left undisturbed during construction. The researchers randomly selected study areas within and outside of the pipeline corridor and then conducted a comparative analysis of species diversity. To calculate the site's species diversity the Shannon 38 Arnoldia 73\/3 Woodland Restoration 39 A study in 2013 assessed plant species in several areas, including this transect across the construction zone. mination within the existing seed bank rather than the introduction of new pioneer species. More importantly, perhaps, was the realization that the seed bank remained viable within the soil blocks that were lifted and reset, and that this habitat restoration method successfully maintained the pre-construction forest, woodland, and wet meadow species profiles while keeping new species at bay. Thirty years later, comparative measurements and observation revealed that the native understory plant communities present in 2013 closely matched the pipeline corridor's preconstruction species. These included mixed- age American beech and understory species like striped prince's pine (Chimaphila maculata), Jack-in-the-pulpit (Arisaema triphyllum), and lady fern (Athyrium filix-femina) (McCoy 2013). This healthy species composition reveals a stark contrast to the nearby Texas Eastern Transmission pipeline, which was constructed around the same time as the Algonquin Gas Transmission pipeline using traditional construction methods. In 2013, the Texas Eastern Transmission pipeline's 100-foot-wide, linear disturbance zone exhibited low species diversity dominated by orchard grass (Dactylis glomerata) and other non-native plants. 40 Arnoldia 73\/3 Woodland Restoration 41 COURTESY OF ROBERT VOLKMAN, JEDROC CONSULTING SERVICES Typical pipeline construction involves clearcutting and environmental disruption of wide swaths of land. This photograph shows installation of a natural gas pipeline in northeastern British Columbia. ties result in human injuries and fatalities as well as environmental damage from spilled hazardous liquids. From 1995 to 2014, 360 fatalities and 1,365 injuries have been associated with gas and hazardous liquid pipelines in the United States, and 2,171,575 spilled barrels of hazardous liquids have been reported to Pipeline and Hazardous Materials Safety Administration (Note that gas leaks are not quantified in PHMSA reports.) These environmental con- cerns are particularly heightened in residential neighborhoods and areas that serve as sources of drinking water. The best way to prevent habitat degradation is, of course, to avoid disturbing sensitive landscapes in the first place. However, when disturbance is unavoidable, minimally-invasive construction techniques paired with ecologically sound restoration practices offer the best possible opportunity for affected ecosys- 42 Arnoldia 73\/3 "},{"has_event_date":0,"type":"arnoldia","title":"A Dandy for Winter: Jasminum nudiflorum","article_sequence":4,"start_page":44,"end_page":45,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25597","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170a36b.jpg","volume":73,"issue_number":3,"year":2016,"series":null,"season":null,"authors":"Hetman, Jon","article_content":"A Dandy for Winter: Jasminum nudiflorum Jon Hetman iscovering eye-catching ornamental interest in the winter landscape can be a challenge, but is by no means an impossible task. Some taxa retain their attractive fruits long into winter, and plants with handsome bark like paperbark maple (Acer griseum) or colorful stems like red osier dogwood (Cornus sericea) stand out like beacons against a snowy backdrop. Truly astonishing, both from a visual standpoint as well as a scientific one, is the odd species that hazards to bloom when most other plants--and indeed most pollinators--lie dormant. Jasminum nudiflorum, or winter jasmine, is one such horticultural jewel. A member of the olive family (Oleaceae), winter jasmine is a hardy member of a genus best known for its potently fragrant tropical and subtropical members. Blossoms of this small deciduous shrub appear before the leaves (its specific epithet means \"naked flowers\"), borne singly in the leaf axils on the previous year's wood like its relative, Forsythia, which it rather resembles. Winter jasmine's small, waxy, bright yellow flowers feature funnel-shaped corollas that flare at the end into five or six spreading lobes, giving a starlike appearance. These are described as either non-fragrant or possessing a delicate, mossy scent, but in any event they do not summon the delightful olfactory sensations that makes its genus name synonymous with perfume. Fortunately this shortcoming is redeemed by an extremely lengthy period of bloom, which may last from November to March. In severe winters, some dieback may occur and flowers may suffer damage, but the plant usually rebounds to continue flowering after such events. Compounding the seasonal interest offered by its flowers, J. nudiflorum also delights with arching, willowy green stems that provide further visual relief from winter's tonal monotony. In spring, stems produce compound leaves that are oppositely arranged and composed of three ovate leaflets, each about "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23447","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170816e.jpg","title":"2016-73-3","volume":73,"issue_number":3,"year":2016,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Seeing the Landscape in Landscape Art","article_sequence":1,"start_page":2,"end_page":18,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25593","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160856f.jpg","volume":73,"issue_number":2,"year":2015,"series":null,"season":null,"authors":"Faison, Edward K.","article_content":"Seeing the Landscape in Landscape Art Edward K. Faison I n 1825, a young British immigrant, captivated by the wild scenery of the Hudson River and nearby Catskill mountains, endeavored to promote America's natural wonders as a distinctive national identity. That year Thomas Cole began painting the undeveloped landscapes of the Northeast with romantic grandeur and literal exactitude, inspiring a cadre of followers that produced America's first painting movement. The Hudson River School (HRS), as the movement was later named, thrived for the next half century before being replaced by the misty, ethereal landscapes of the tonalists COURTESY OF THE METROPOLITAN MUSEUM OF ART made famous by George Inness in the 1880s and 1890s. In an ironic twist, a painting fraternity (the HRS) founded to celebrate America's wilderness became synchronous with a brief period in the northeastern United States in which the landscape was altered to a greater extent than at any time since the last ice age. Because photography was in its infancy during this period and because intensive observation and faithful depiction of nature as well as the study of natural science were integral to the HRS's ethos, nineteenth century American landscape painting affords a window into the dramatic ecologi- Thomas Cole's 1836 painting, View from Mount Holyoke, Northampton, Massachusetts, after a Thunderstorm-- The Oxbow. Cole included a portrait of himself working at his easel, dwarfed by the surrounding forest, in the lower center of the painting. Landscape Art 3 COURTESY OF THE NATIONAL GALLERY OF ART, WASHINGTON, D.C. Deforestation is evident in George Inness's The Lackawanna Valley, circa 1856. cal changes that occurred across the region. In turn, these spectacularly rendered landscapes, when viewed with an eye toward ecology and natural history, can be seen afresh. From Forests to Fields Few paintings capture the overarching landscape dynamic of nineteenth century northeastern North America as effectively as Cole's View from Mount Holyoke, Northampton, Massachusetts, after a Thunderstorm--The Oxbow. Painted in 1836, The Oxbow depicts a wild, storm-battered forest clinging to the slopes of Mount Holyoke under a darkened sky, juxtaposed against a sunlit, cultivated landscape surrounding the Connecticut River's oxbow below. Cole seems to capture the moment just before humanity on the right sweeps across the canvas and conquers the remaining wild nature on the left. Cole was certainly aware of and somewhat ambivalent toward the dramatic changes to the land that were occurring around him. Although he admired the cultural achievements of Europe and anticipated similar cultural greatness in America, he also decried the rapid loss of forest that inevitably accompanied the advancement of civilization. In 1841, Cole wrote on behalf of the forest: Our doom is near ... These slumbering mountains, resting in our arms, Shall naked glare beneath the scorching sun, And all their wimpling rivulets be dry. No more the deer shall haunt these bosky glens, Nor the pert squirrel chatter near his store. A few short years! --our ancient race shall be, Like Israel's, scattered `mong the tribes of men. Cole wasn't far from the truth. In fact he was witnessing one of the greatest acts of deforestation the world has ever known. While forest clearance took several centuries in Europe, in eastern North America it was largely condensed into two generations. From about 1810 4 Arnoldia 73\/2 Landscape Art 5 Jasper Cropsey's The Valley of Wyoming, 1865, depicts the agrarian landscape that replaced eastern forests. Courtesy of the Metropolitan Museum of Art. everywhere except for a few inhospitable and temporarily disturbed locations. As forest ecologist E. Lucy Braun (1950) wrote: When the Pilgrims came to this continent, New England was covered by forest interrupted only where lakes or bogs and river swamps made tree growth impossible; where sand deposits near the coast were unsuitable for closed stands; where fire or windfall had temporarily destroyed the forest; where Indians had burned the forest (especially near the coast); and where rock outcrops occurred in the more rugged sections. cousins of modern day elephants, were common in this transitional landscape between tundra and forest, and these large herbivores probably helped maintain the landscape's semiopen character, much the way elephants do in African savannas today. Disturbances in the Nineteenth Century Landscape The tranquility evoked by Cropsey's Valley of Wyoming belies the relentless disturbances required to maintain agrarian landscapes of the Northeast in a semi-open state. However, a closer look at the composition reveals some of these disturbances. In the left middle ground, we see farmhands cutting and collecting hay in an upland meadow near a gray barn. The arduous task of cutting hay meadows by hand provided fodder for livestock in winter, and simultaneously prevented trees and shrubs from invading and overtaking the grass. Cattle One would have to travel back 12,000 to 14,000 years to the end of the last ice age to find an environment that supported open landscapes in the Northeast at a scale comparable to the agrarian landscapes of the nineteenth century. Then, cold climates south of the waning ice sheet sustained a mix of tundra grasses and sedges and scattered spruce trees in an open \"spruce parkland.\" Mastodons, the now extinct 6 Arnoldia 73\/2 Landscape Art 7 forbs (\"wildflowers\"). Understory shrubs and trees are sparse, and the ground flora is essentially a continuation of the adjacent meadow--a vegetation structure pleasant enough for a late afternoon stroll by the woman in the center of the composition. Inness's \"forest\" would be described by ecologists today as a savanna or open woodland. His title and composition therefore reveal much about the structure and disturbances of nineteenth century woodlands near settlements. By removing smaller woody plants, burning and grazing often left mature and fire resistant trees (e.g., oaks) to grow larger with reduced competition. Fire and grazing also reduced or removed the leaf litter, releasing herbaceous plants from the suppressive cover of the dead leaves. Selective cutting of trees for fuelwood further increased the openness of these stands, casting more light on the forest floor and promoting a thriving herbaceous layer. The tall wildflowers emerging above the grasses in the foreground create both depth and balance in the composition and are consistent with the effects of cattle preferentially grazing grasses over forbs. Large Wildlife--Rare Symbols of the Wilderness In 1856 Henry Thoreau lamented the depauperate large wildlife community in the fields and ALLEN MEMORIAL ART MUSEUM, OBERLIN COLLEGE. GIFT OF CHARLES F. OLNEY. BRIDGEMAN IMAGES The white-tailed deer in Thomas Cole's 1825 landscape painting Lake with Dead Trees symbolize untamed wilderness. 8 Arnoldia 73\/2 Landscape Art 9 THE HAGGIN MUSEUM, STOCKTON CA Albert Bierstadt's Moose was painted sometime after 1880 from sketches he made along the Maine\/Nova Scotia border, an area that was one of the last strongholds in the Northeast for the species. 10 Arnoldia 73\/2 Landscape Art 11 the HRS's celebration of the American wilderness was getting started. Few other HRS paintings depict wolves or other top predators such as mountain lions, wolverines, and black bears, all of which were hunted with similar fervor and suffered precipitous declines in the mid to late nineteenth century (all except black bears were completely extirpated from the region by the end of the century). Remnant Old Forests Despite the widespread transformation of forests to fields, as well as the dramatic alteration of farm woodlands, relatively sizeable tracts of old growth forest still existed in the mid-nineteenth century Northeast (see Greeley virgin forest map 1850, on page 14). Asher Durand demonstrated a strong affinity for painting undisturbed forest compositions and espoused a particularly strong ethos for representing nature truthfully, stating: \"never let [the art- ist] profane [nature's] sacredness from a willful departure from the truth ... For I maintain that all art is unworthy and vicious that is at variance with truth.\" In Adirondack Mountains, N.Y. (circa 1870), Durand reveals an extensive forested plain of seemingly undisturbed wilderness with a weathered hardwood and hemlock standing sentinel-like on a cliff in the right foreground. The closer hardwood has few largediameter limbs in the crown and a relatively small leaf area to trunk volume, suggesting a very old tree. As depicted in Greeley's forest maps, the Adirondacks in northern New York were, indeed, one of the remnant strongholds of old growth forest in the Northeast in the late nineteenth century. Durand also takes us into a forest interior in Forest in the Morning Light (1855). Bryophytes grow high on the trunks of hardwood trees, and moss covers the forest floor, which is strewn with multiple pieces of large downed NEW YORK HISTORICAL SOCIETY. LUCE CENTER 1932.10 Asher Durand's Adirondack Mountains, N.Y., circa 1870, depicts a large expanse of undisturbed forest. 12 Arnoldia 73\/2 COURTESY OF THE NATIONAL GALLERY OF ART, WASHINGTON, D.C. Landscape Art 13 wood. Trees range in size and age from sapling to large veteran. The tree leaning to the right has a low taper (i.e., little difference in diameter) from the base of the trunk to the base of the crown. All of these attributes suggest old age and are characteristic of old growth forests in the Northeast. Interestingly, the species Durand chose to include in this particular composition--an apparent white oak (Quercus alba) leaning to the right in the foreground, an American beech (Fagus grandifolia) with smooth gray bark to the right of the white oak, and perhaps an eastern hemlock (Tsuga canadensis) or white pine (Pinus strobus) in the left foreground--were dominant species of the forests that greeted the first European settlers. Beech was the undisputed king of northern New England, northern New York, and northern Pennsylvania forests, with hemlock the second most important tree. White oak dominated the forests of the southern half of the region. EDWARD K. FAISON This 2015 photograph from Mount Holyoke showing the Connecticut River oxbow was made from approximately the same location that Thomas Cole painted The Oxbow in 1836. 14 Arnoldia 73\/2 Landscape Art 15 sharply in the second half of the twentieth century, expanding outward from nineteenthcentury refugia and thriving on nuts and acorns in maturing beech and oak forests, as well as on readily available deer fawns. A new wild canid, the coyote, migrated into the region from the western plains, partially filling the vacated niche left by the extirpated wolf. In its eastward expansion, the coyote interbred with wolves in the eastern Canadian provinces, producing a larger version of its western progenitor and an animal capable of bringing down deer. More recently, the vanguards of extirpated large carnivores have begun passing through the Northeast. At least four wild gray wolves and four wild cougars have been confirmed in the region in the past two decades, and unconfirmed sightings of cougars have increased dramatically. Of course, not everything has returned to a wilder condition today compared with 1836. Gone is an avian wonder that Cole may have seen from Mt Holyoke's summit: the passenger pigeon. This species once congregated in flocks in the millions, even billions, before being robbed of its forested habitat and hunted to extinction by the end of the nineteenth century. Wolverines still occurred in the Adirondacks as of 1842 and were reported to be in Pennsylvania, Maine, Vermont, and even in the Hoosac range of Massachusetts in the nineteenth century. These largest members of the weasel family remain far north of the United States today in upper Quebec and Newfoundland. Elk still roamed parts of New York and caribou inhabited northern Maine in the midnineteenth century, but both animals remain extirpated from those states today. There are also far fewer old growth forests today than in 1836, even if the percentage of forest area today is higher. Forest greater than 200 years in age cover only about 0.4% of the northeastern United States, compared with the relatively sizeable tracts of old growth forest in the mid-nineteenth century (see 1850 Greeley map, on facing page). The long-lived beech, white oak, and hemlock that dominated early colonial forest composition, have been replaced by shorter-lived and earlier successional species such as red maple, black cherry (Prunus serotina), and birches in the younger forests that have grown back on abandoned farmland and cutover lands. The large pieces of downed wood and moss covered trees in Durand's Forest in the Morning Light are far less common in today's drier, second growth forests. Forest wildflowers are generally less abundant and diverse in second growth forests compared with old growth forests, and bird densities are also lower in the former compared to the latter. Novel Threats One of the factors that has slowed the recovery of beech and hemlock to their former positions of dominance in northeastern forests is the invasion of forest pests and pathogens from Eurasia. The exotic fungus Nectria coccinea var. faginata, introduced to Nova Scotia in the early 1900s, has subsequently spread throughout the Northeast, invading the bark and killing many mature beech trees. Hemlock woolly adelgid, an aphidlike insect introduced from Japan, reached New England in 1985 and has thinned the canopy and killed many hemlocks in the southern parts of the Northeast. But by far the most dramatic change to the modern forest resulting from an introduced forest pest is a tree portrayed in William Trost Richards's October (1863). Richards was a member of a brief movement in the 1850s and 1860s that called themselves the Association for the Advancement of Truth in Art. Inspired by the British art critic John Ruskin, the American Pre-Raphaelites (as the group was later named) took the accurate portrayal of nature to a new level. In October, Richards's highly detailed rendering of an autumn forest scene enables us to identify the large tree on the right of the composition with diamond-shaped furrows on the trunk and linear sawtooth leaves with considerable confidence: an American chestnut (Castanea dentata). A chestnut this size hasn't been seen in the forests of the Northeast in perhaps 75 years. Forty years after Richards's painting, the Asian chestnut blight (Cryphonectria parasitica) arrived in New York City, and over the next several decades destroyed virtually every mature chestnut throughout its Appalachian 16 Arnoldia 73\/2 COURTESY OF THE NATIONAL GALLERY OF ART, WASHINGTON, D.C. Landscape Art 17 mountain range. Chestnuts today rarely grow above 15 centimeters (6 inches) in diameter, perhaps 1\/5 to 1\/6 the diameter of the tree in Richards's painting, before being killed by the blight and therefore almost never emerge into the forest canopy. Interestingly, the smaller tree to the left of the chestnut in the right foreground, flowering dogwood (Benthamidia florida; syn. Cornus florida), has also been struck by an exotic fungus, dogwood anthracnose (Discula destructiva), and has declined significantly in recent decades. The Future Landscape: Stemming a Second Deforestation Looking north from the summit of Mount Holyoke, past the farm fields that have succeeded to forest, an unmistakable threat to the forested landscape can be seen: the patchwork of residential and industrial development and roads fragmenting and perforating the forests and farm fields. As the forest grew back in the twentieth century after farm abandonment, human populations also surged. By 1975, the human juggernaut caught up to the regenerating forests, and the pendulum of 100 years of forest recovery began to swing back towards forest loss (see forest and population map). Forty years later, forest loss is in near free fall in New Hampshire and the southern New England states. More recently, Vermont began losing forest at an increasing rate. Only Maine has been able to sustain a balance between forest loss and forest recovery, although residential development is projected to increase significantly in the southern part of the state over the next two decades. This deforestation is much harder for trees to recover from than before. Paved roads and housing developments represent a \"hard deforestation,\" in contrast to the \"soft\" deforestation of agricultural fields in the nineteenth century (Foster et al. 2010). How will the northeastern landscape look in the next 50 to 100 years? The answer depends in large part on whether conservation groups, private landowners, public agencies, and other stakeholders are willing to work together to protect both forest and farmland. The news so far is promising. In the past 10 to 15 years, part- nerships of conservation groups that transcend political boundaries have increased by a factor of six in New England and adjacent New York. This type of regional collaboration is at the heart of The Wildlands and Woodlands Vision created by 20 scientists and environmentalists across the region. The Vision calls for the permanent protection of 70% (30 million acres) of the New England region in forest over the next 50 years. Like the structure of an ecosystem, the Wildlands and Woodlands Vision is fundamentally a bottom up (grass roots) effort. It has to be: over 80% of New England's forestland is privately owned. If he were alive today, Thomas Cole would be amazed to see more forest cover in New England than he saw in 1836. But he would once again recognize and lament the signs of deforestation. History has inadvertently given us a second chance to live in a forested New England, but there will be nothing inadvertent about the efforts needed to keep these forests standing. References Askins, R. A. 2000. Restoring North America's Birds. New Haven: Yale University Press. Askins, R. A. 2014. Saving the World's Deciduous Forests. New Haven: Yale University Press Aubry, K. B., K. S. McKelvey, and J. P. Copeland. 2007. Distribution and broad-scale habitat relations of the wolverine in the contiguous United States. Journal of Wildlife Management 71: 2147 18 Arnoldia 73\/2 "},{"has_event_date":0,"type":"arnoldia","title":"The Etymology of Parking","article_sequence":2,"start_page":19,"end_page":24,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25595","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160896d.jpg","volume":73,"issue_number":2,"year":2015,"series":null,"season":null,"authors":"Richmond, Michele","article_content":"The Etymology of Parking Michele Richmond park (circa 1845) verb a. to plant a tree or spread a patch of turf or flowers b. to create a little patch of parkland park (circa 2015) verb a. (1) to bring a vehicle to a stop and keep standing at the edge of a public way (2) to leave temporarily on a public way or in a parking lot or garage b. to enclose in a park I 've always wondered why we use the word parking to describe a place to leave a car. For me the word evokes images of my neighborhood park, playgrounds, or New York's Central Park: lush green spaces, not places easily reconciled with a patch of asphalt. A few years ago while I was working at the New York City Department of Parks and Recreation, I finally got my answer. While exploring the history of street trees, I came upon a law passed by the United States Congress on April 6, 1870, authorizing the city SHAWN STANKEWICH Public green spaces like New York City's Central Park, seen here, add to the livability of urban areas. 20 Arnoldia 73\/2 DUMBARTON OAKS Etymology of Parking 21 Automobiles Arrive The world was changing rapidly as the twentieth century arrived. The number of automobiles in the United States increased from 8,000 in 1900 to over 8 million in 1920 and marked a major shift in the meaning of the term parking. Just as people would tie their horses to the parking trees, automobiles began to stop next to the parking strips lining each road. The increase in the number of automobiles on the road, the enhancements made to the National Mall, and the See America First tourist campaign, which began in 1910, led to a huge increase in the number of cars in Washington, D.C., from both locals and tourists. The See America First campaign was designed and implemented by America's railroad companies and advertised America's first National Parks. Advertising for the National Parks, the majority of which were located in the western United States in the early 1900s, benefited the railroads immensely: more tourists journeying out west meant more money. However, the railroads did not foresee the rapid growth of the automobile and an unintended consequence of the See America First campaign was regional tourism (as opposed to national tourism). With all the positive press Washington, D.C., received from the greening of their streets through parking, the city received more This image from a 1915 National Geographic article about Washington, D.C., shows both an early automobile and a horsedrawn carriage taking advantage of shade from street trees. tourists than any other city in America in the 1920s and 1930s. Of course, the Washington, D.C., Parking Commission had not planned for the automobile when setting out their parking system. By the mid-1920s city officials began cutting down street trees and widening streets to accommodate the volume of cars, thereby replacing the original meaning of parking as a place for trees and greenery with parking as a place for automobiles to stop. Some of the earliest instances of this shift appear in Washington Post articles from the 1920s, where the term \"parking\" was used to explain where cars were parked rather than to where trees were planted. BIODIVERSITY HERITAGE LIBRARY place wooden boxes around the trees to protect them, but with the passage of another congressional law placing the jurisdiction of parking places squarely in the hands of the Commissioners of Washington, D.C., this practice was soon discarded. The new law had unintended consequences: the removal of the protective boxes allowed people to wedge their way into the parking system. How? Because during the hot summers in Washington, trees provided shade for horses while their owners were off in a shop or visiting a friend. Owners would tie their horses (and carriages) up to the street trees, effectively decreasing a two-lane road to one active lane and one stopped lane. Although it became illegal in 1882 to trespass on parking, or to cut, injure, or maim parking trees in any way, the convenience and shade provided by the trees for the waiting carriages and horses outweighed the fine levied. An illustration from the article \"The New Washington\" (The Century Magazine, March 1884) shows a protective structure around the trunk of a street tree on Thomas Circle in Washington, D.C. COURTESY OF HATHITRUST 22 Arnoldia 73\/2 LIBRARY OF CONGRESS Etymology of Parking 23 MICHELE RICHMOND Extensive plantings and wide sidewalks make city streets more usable and inviting. dents to determine what they wanted on their street. Since car parking spaces are useless to many low-income San Franciscans, the neighborhood ranked street amenities (from highest priority to lowest priority) as: walking, street trees, public transit, biking, and car parking. These rankings have guided the redesign of the street away from parking towards a multimodal street with a significant increase in the number of planned street trees. In another shift, Seattle has prioritized the creation of Green Streets throughout the city. Seattle Department of Transportation's U District Green Streets Concept Plan emphasizes the planting of healthy, consistently placed street trees, calling street trees crucial to the character and livability of the city. Their plan prioritizes street trees, perennial and shrub plantings, bike parking, and pedestrian movement over car parking along the Green Streets. The City of Toronto has set a goal of increasing their overall tree canopy of 10.2 million trees by approximately 13 percent over the next few decades. The city recognizes the wide range of environmental, ecological, social, cultural, and economic benefits their urban forest provides for residents and has developed multiple strategies to increase their overall tree canopy. At the street level, the city has worked diligently to increase street tree planting and survival rate in conjunction with better public transit and ease of walking. One strategy they have implemented is the use of below pavement soil structural systems (such as Silva Cells) that allow soil to receive air and water without being compacted by people, bikes, and buses moving above them. The strategy has been so successful that Silva Cells are now being deployed across the city to allow large shade trees to grow successfully in congested urban conditions. The story of parking over the past 140 years exemplifies changing social norms, ways in which city planners absorb technological advancements such as the birth of the automo- 24 Arnoldia 73\/2 "},{"has_event_date":0,"type":"arnoldia","title":"The Bonsai Collection of Stellenbosch University Botanical Garden","article_sequence":3,"start_page":25,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25594","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d1608928.jpg","volume":73,"issue_number":2,"year":2015,"series":null,"season":null,"authors":"Pretorius, Willem; Sax, Miles S.","article_content":"The Bonsai Collection of Stellenbosch University Botanical Garden Miles S. Sax and Willem Pretorius A panoramic view of Stellenbosch, Western Cape, South Africa. FMALAN AT EN.WIKIPEDIA S outh Africa has long been recognized for its picturesque landscapes, large wildlife species, and stunning plant diversity. Housing roughly 10 percent of the world's vascular plant biodiversity (about 30,000 taxa) on less than 1 percent of the earth land surface, it is the only country in the world that contains its own floristic kingdom (the Cape Floral Kingdom) within the confines of its borders. With plant endemism rates as high as 70 percent in the Western Cape, South Africa has continued to fascinate plant explorers, botanists, and horticulturists. Flowering geraniums (Pelargonium spp.), bird-of-paradise (Strelitzia reginae), and the calla lily (Zantedeschia aethiopica) are just a few of the South African plants that are now known to gardeners worldwide. A Garden Grows in Stellenbosch The town of Stellenbosch is located in South Africa's Western Cape province. It lies on the eastern flank of the Cape Flats, situated at the base of the commanding Hottentots-Holland Mountains, which provide an inspiring back- drop to the municipality. The Stellenbosch University (originally called the Stellenbosch Gymnasium) was established in this agricultural and wine-producing region in 1866. In 1902, the University made efforts to establish the first botanical garden in Africa to be housed in an institute of higher education. Augusta Vera Duthie was the first lecturer in botany at Stellenbosch and undertook this initiative in order to grow plant material for educating botany students at the school. In 1902, the first garden was laid out in front of the Main Building where it remained for twenty years. In 1922, under the guidance of Dr. Gert Cornelius Nel, the garden was moved to accommodate the growing collections and to establish a permanent site. The 1922 design included transectional and circular order beds for botanical education and were inspired by the world's oldest botanical garden in Padua, Italy. As Stellenbosch University Botanical Garden (SUBG) has developed over the decades its plantings and design have changed in response to the evolving mission of the garden and the priorities set 26 Arnoldia 73\/2 Bonsai Collection of SUBG 27 MILES S. SAX Clockwise from upper left: Narrow-leafed bird-of-paradise (Strelitzia juncea); A formal pond at Stellenbosch University Botanical Garden displays water lilies, including the platter-like Santa Cruz water lily (Victoria cruziana); Drosanthemum bicolor, a South African succulent with showy flowers. A Unique Bonsai Collection One collection in particular at SUBG stands out for its distinctive character--the bonsai collection, whose plants have played a central role in the development of a unique horticulture practice in Africa. Known as the Western Cape Heritage Bonsai Collection, it is the oldest and largest public bonsai garden in Africa. What makes this collection special is the wide variety of indigenous species used, its development by a cast of interesting characters, and the display of the internationally recognized African styles of bonsais. Originally established in 1972, the bonsai collection was incorporated into the gardens by Wim Tijmens, SUBG curator from 1962 to 1999. Wim is recognized for establishing much of the SUBG layout and design that provide the defining elements of the landscape today. His passion for the temperate flora of East Asia took 28 Arnoldia 73\/2 Bonsai Collection of SUBG 29 Louis Nel with his outstanding Buddleja saligna bonsai trained in the Pierneef style. tion and instruction of bonsai in South Africa. When she donated her collection, SUBG curator Wim Tijmens had some trouble explaining the significance of the gift to the University's chancellor, who didn't know what bonsai was. The next big donation to the collection came from the Reverend Gerjo van der Merwe, a minister in the Dutch Reformed Church. According to his family, he was one of the first bonsai enthusiasts to grow indigenous bonsai from seeds in South Africa. From Van der Merwe's personal notes his deep love of nature and the belief in the healing power of trees shines through as an obvious motivator for his bonsai passion. Because he was often transferred, he decided to use soil from the family farm in Boplaas in which he transported small trees with him on his travels. \"I believe that the growing of bonsai is a healthy help to heal this broken relationship. By growing trees from your own region and to take it with you is a strong way to keep and protect the association with a specific place and the sense of belonging.\" The third significant donation of bonsai to the collection was that of Louis Nel. His collection was donated on March 20, 2012. Nel was internationally known as the king of Buddleja saligna, an evergreen South African species commonly known as false olive or squarestem butterflybush. Nel started working with bonsai in 1974 and his skill and reputation quickly grew. Throughout his life he was a regular contributor to bonsai magazines and participated in many international bonsai demonstrations. A number of his trees won international competitions; an outstanding example is SUBG accession 2012-1, one of Nel's Buddleja saligna bonsai. This tree now stands as one of the garden's premier specimens, highlighting a South African native plant as well as a style of SUBG 30 Arnoldia 73\/2 Bonsai Collection of SUBG 31 SUBG it trend towards the less exacting and playful penjing style of China. The tree species found in the SUBG collection weave their own story of the development of bonsai in South Africa. The oldest trees in the collection date back to the 1940s and were created by South African bonsai pioneer Becky Lucas. Her original trees are the classic bonsai subject, Japanese black pine (Pinus thunbergii), which is native to Japan, China, and Korea. As Lucas continued to develop her passion and skill in bonsai, she mostly used the traditional Eastern pallet of plant species including trident maple (Acer buergerianum), deodar cedar (Cedrus deodara), and Atlas cedar (Cedrus atlantica). As the SUBG An African olive (Olea europaea subsp. africana, syn. O. e. subsp. cuspidata) collection developed over the bonsai created by Louis Nel. decades and new bonsai growers a popular choice for growers for many centuries. added their contributions, many other familiar For example, a specimen of Ficus retusa (syn. bonsai subjects joined the collection including dwarf Japanese garden juniper (Juniperus F. microcarpa) in the Crespi Bonsai Museum procumbens `Nana'), lacebark or Chinese elm in Italy is purported to be over 1,000 years old. (Ulmus parvifolia), and Ligustrum ovalifolium, Ficus natalensis was likely selected to be trialed in bonsai culture as a substitute for East which oddly goes by the common name California privet despite being native to Japan and Asian species because it has traits that emulate South Korea. those of traditional bonsai species. Over the Although South African bonsai started out years Natal fig has proved itself as an excellent selection for development as bonsai and with traditional Eastern influences it has its popularity has expanded, making it a mainbecome adapted and transformed as a result stay for growers in South Africa and abroad. F. of its interpretation in Africa. It is unclear natalensis makes up one of the largest groups what prompted the beginning of use of African plant species for bonsai in South Africa. of species in the SUBG collection, comprising It can be speculated that perhaps a lack of around 10 percent. suitable imported plant material from Japan Another African species that has become and East Asia during World War II might have very popular for bonsai is Olea europaea subsp. limited availability. africana (syn. O. e. subsp. cuspidata), a subordinate taxon of the commonly known European The first African tree in the collection to olive tree. The distribution of this subspecies be adapted for bonsai was the Natal fig (Ficus ranges widely from Africa through the Middle natalensis) by Ian Ross in 1948. The native East, India, and China. It is widely grown in range of Ficus natalensis extends from Mozambique, Malawi, and Zimbabwe to South Africa. parks and urban environments for its stress tolerance and cold hardiness (Joffe 2002). The speIt is cultivated in southeastern Africa as a shade cies displays small, lustrous, evergreen leaves tree and living fence. The genus Ficus is commonly used for traditional bonsai and has been that form dense canopies, and greenish white Popular South African Styles Of the six styles described by Charles Ceronio, the three that have gained widespread adoption in South Africa are the flattop, Pierneef, and baobab styles (Adam 1992, Ceronio 1999). Baobab style This style focuses on mimicking the iconic African baobab tree (Adansonia digitata). Baobabs in the wild are considered to be one of the world's largest trees, not for their height, but for their impressive girth. These trees tend to grow with massive central trunks giving away to relatively short branches. Adapted to growing in regions with high summer temperatures, the genus is found throughout tropical Africa and Madagascar. For the baobab bonsai style the focus is placed on the upright trunk habit, giving the impression of a large central section while the branches are kept short. Seldom seen in collections, this style can be accomplished by using taxa such as the African baobab itself, corkwood (Commiphora spp.), or coral tree (Erythrina spp.). In order to convincingly emulate the thick trunk, bonsai trainers often select several upright stems and use twine or wire to pull the multiple stems together, thus fusing them into a single trunk. This style was principally developed by Charles Ceronio. A baobab (Adansonia digitata) growing in South Africa's Kruger National Park. REBECCA PRETORIUS A 28-year-old baobab bonsai created by article co-author Willem Pretorius. MILES S. SAX Pierneef style The Pierneef style is an open umbrella-shaped tree form. It invites the viewer to imagine the tree spreading wide over an expansive savanna landscape. The umbrella form itself is created by the tight zigzag branching pattern reminiscent of acacia species, such as Acacia sieberiana (syn. Vachellia sieberiana), found throughout Africa. The style was named after South African artist Jacobus Hendrik Pierneef (1886 34 Arnoldia 73\/2 Bonsai Collection of SUBG 35 SUBG Many plant species and bonsai styles are on display in Stellenbosch University Botanical Garden's bonsai house. there is some question as to the longevity of this tree for bonsai because of the shrub's soft wood. The collection of Buddleja specimens at SUBG will stand as a long term trial of the species' suitability as a bonsai subject. Just over 10 percent (23 of 213 specimens) of the collection is made up of this single taxon. Bibliography Adam, R. 1992. Bonsai in South Africa. Cape Town: Struik Publishers Ltd. Ceronio, C. 1999. Bonsai Styles of The World. Pretoria: Charles S. Ceronio. Hemy, C. 1967. How to Grow Miniature Trees, Full instructions for practising the ancient art of bonsai. Supplement to Farmer's Weekly, January 25. Joffe, P. 2002. PlantZAfrica. http:\/\/www.plantzafrica. com\/frames\/plantsfram.htm (accessed January 22, 2015). Lucas, B. 1968. A Summer in Japan. Bonsai Magazine 7(3): 8"},{"has_event_date":0,"type":"arnoldia","title":"Tracking the Seasonal Rhythms of Boston Common Trees","article_sequence":4,"start_page":36,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25596","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d170a326.jpg","volume":73,"issue_number":2,"year":2015,"series":null,"season":null,"authors":"Richardson, Andrew D.; Oswald, W. Wyatt","article_content":"Tracking the Seasonal Rhythms of Boston Common Trees W. Wyatt Oswald and Andrew D. Richardson N ew England's deciduous forests undergo dramatic seasonal changes. New leaves emerge from protective buds as winter gives way to spring, green trees conceal the landscape and cast deep shade during the height of summer, and glorious fall foliage senesces as autumn yields to the snows and bitter cold of winter. But how much do these seasonal transitions vary from year to year? We are exploring this phenological question through detailed observation of a setting familiar to many New Englanders and visitors alike--the Common in downtown Boston. Since mid May of 2010, a digital camera mounted on the roof of the ten-story Walker Building, overlooking the Boston Common from the campus of Emerson College, has taken photos at thirty-minute intervals of a tree-covered area on the east side of this historic park. The images generated by digital cameras represent colors using the RGB (red, green, blue) additive color model, which proposes that any color perceived by the human eye can be represented by some combination of these three primary colors. Each digital image is actually composed of three separate layers, one each for red, green, and blue. We characterize the \"greenness\" of the tree canopy by using image analysis software to measure how bright the green layer is relative to the total brightness of the red, green, and blue layers together. The color of the canopy is related to the color of individual leaves, as well as to the number of layers of leaves in the canopy. Individual leaf color is largely determined by pigments--green Photos of the Boston Common in spring (April 30, 2011), summer (July 1, 2011), fall (November 11, 2011), and winter (February 5, 2012). ALL IMAGES BY THE AUTHORS UNLESS OTHERWISE NOTED Tracking the Seasonal Rhythms 37 chlorophylls, orange carotenoids, and red anthocyanins--but is also an indicator of photosynthetic capacity (Richardson et al. 2007; Sonnentag et al. 2012). When we use this approach to quantify the greenness of all of these photos of the Common we can visualize the seasonal shifts. Following budburst, leaves expand rapidly and the springtime green-up happens quickly. Over a span of just four weeks, the leaf-out of deciduous elms, basswoods, cherries, and maples transforms the Common from a late winter landscape of browns and grays to its maximum greenness, which generally occurs during the first half of May. Peak green lasts only a couple of weeks, though, because as leaves mature they actually darken somewhat (Keenan et al., 2014). This causes a gradual reduction in our greenness index over the course of the summer. Then fall arrives: day lengths get shorter, temperatures become colder, and chlorophyll production gradually slows down. Greenness fades with the onset of senescence, leaf coloration, and leaf fall. The timing of these autumnal changes varies from species to species, and thus the de-greening of the Boston Common landscape happens somewhat more slowly than spring leaf-out, taking about six weeks from start to finish. Comparison of the Boston Common data with those from a similar camera at Harvard Forest, located in the north-central Massachusetts town of Petersham, illustrates the general similarity of vegetation phenology in deciduous forests across southern New England. Despite differences in species composition between the native flora of Harvard Forest and the humanconstructed mixture of native and nonnative tree species that we find in the Boston Common, both landscapes feature rapid spring greenup and maximum greenness at the beginning of the growing season. However, since Harvard Forest is situated more than 300 meters (984 feet) higher than Boston, nearly 100 kilometers (62 miles) inland, and well outside the urban heat island (Zhang et al. 2004), it is cooler than the Common and thus has a shorter growing About Boston Common Since its founding in 1634, the fifty-acre Boston Common has served many purposes. In the seventeenth century, the sparsely wooded Common was used as a pasture for cattle. Early maps show only three trees, including the Great Elm, a majestic tree that loomed over the Common until it was blown down in a windstorm in the winter of 1876. The Common began to take on parklike qualities early in the eighteenth century. Bostonians strolled along wide, tree-lined malls, the first of which was established along Tremont Street in 1722. During the American Revolution and War of 1812, however, hundreds of soldiers were based in Boston and large encampments were built on the Common. Cows were eventually banned in 1830, and with subsequent formal landscaping the Boston Common was transformed to the largely forested park that exists today (Friends of the Public Garden 2005). This 1845 engraving by Hammatt Billings shows the Great Elm growing in the Boston Common. COURTESY OF HISTORIC NEW ENGLAND 38 Arnoldia 73\/2 Tracking the Seasonal Rhythms 39 season. With earlier leaf-out and later leaf drop, trees in downtown Boston have green leaves some eight weeks longer than their counterparts in Petersham. Now that we have captured the vegetation phenology of five springs in the Boston Common photos, we can see how the timing of leaf-out has varied across years with strikingly different weather conditions. All but one of the years have a similar pattern: in 2011, 2013, 2014, and 2015, leaf emergence at the Common started at the beginning of April and greenness increased steadily over the next five weeks or so. In contrast, early spring temperatures in 2012 were significantly warmer than normal (Friedl et al. 2014), with record-breaking heat in Boston during the middle of March. The trees of the Boston Common responded immediately to this unseasonable warmth, leafing out within just a few days. As a result, the 2012 growing season was about a month longer than the other years for which we have photos. The first five years of data from the Boston Common are consistent with other phenology studies in showing that the green-up of deciduous trees is highly sensitive to climatic variability (Cleland et al. 2007). If the warmth of March 2012 gives us a sense of what the future holds for springtime in Boston, as climate projections suggest (Hayhoe et al. 2007), then we can expect earlier leaf-outs and a substantial lengthening of the growing season. A failure to track these changes could be deleterious to the insect, bird, and mammal species that utilize urban forests as habitat. Such an outcome also has consequences beyond the Boston Common: phenological changes have the potential to shift competitive interactions among tree species and affect the carbon balance of forest ecosystems (Keenan et al. 2014). To better anticipate these dynamics, our research on vegetation phenology and related ecological processes will continue for years to come at the Boston Common, Harvard Forest, and dozens of other sites where this type of study is being carried out as part of the PhenoCam Network (phenocam. sr.unh.edu\/webcam). More broadly, as Aldo Leopold put it, \"phenology may eventually shed some light on that ultimate enigma, the land's inner workings\" (Leopold and Jones 1947). References Cleland, E. E., I. Chuine, A. Menzel, H. A. Mooney, and M. D. Schwartz. 2007. Shifting plant phenology in response to global change. Trends in Ecology and Evolution 22: 357"},{"has_event_date":0,"type":"arnoldia","title":"A Study in Scarlet: Nyssa sinensis","article_sequence":5,"start_page":40,"end_page":40,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25592","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160816b.jpg","volume":73,"issue_number":2,"year":2015,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"A Study in Scarlet: Nyssa sinensis Nancy Rose M y favorite \"old reliables\" for fall color at the Arboretum include the pure gold foliage of sweet birch (Betula lenta), the fiery red-orange-yellow display of Korean maple (Acer pseudosieboldianum), and the glossy burgundy leaves of Euonymus carnosus. That's just a start, though, and one of the delights of wandering the Arboretum repeatedly in autumn is discovering new spots of color. A few years ago, on a gray mid-November day when many trees were already bare, I was drawn to a cluster of brilliant scarlet and orange leaves remaining on a tree branch. The tree, it turned out, was Chinese tupelo, Nyssa sinensis. This was a new species to me, but I certainly knew another species in the genus, Nyssa sylvatica, known by the common names sour gum, black gum, tupelo, black tupelo, pepperidge, or, to residents of Martha's Vineyard, beetlebung. Under any name, this eastern North American species is a handsome tree, noted for its lustrous green foliage that turns stunning shades of red in the fall. Though its native range is thousands of miles away, Chinese tupelo is strikingly similar to sour gum, providing yet another interesting example of the disjunct floras of eastern North America and eastern Asia. Chinese tupelo is slightly smaller than sour gum, growing as tall as 60 feet (18 meters) in the wild but in cultivation typically reaching only 30 to 40 feet (9 to 12 meters). It has a pyramidal to upright-oval form and dark gray, irregularly fissured bark. Chinese tupelo is deciduous; when its leaves emerge in late spring they often have an attractive red or bronze tint. Mature leaves are about 4 to 6 inches (10 to 15 centimeters) long, dark green and slightly shiny above, lighter green below. Autumn foliage color may be variable depending on individual plant and local climate, but typically ranges from light yellow and apricot to bright, almost luminous reds and oranges. As with other Nyssa species, Chinese tupelo is primarily dioecious (male and female flowers borne on separate plants) but some plants may also bear a few perfect (having both male and female parts) flowers. Female flowers are borne in axillary clusters and male flowers are produced along older branches. The small greenish flowers are inconspicuous but they are extremely attractive to honeybees (N. ogeche, which has a limited native range primarily in southern Georgia and northern Florida, is the source for prized tupelo honey). The fruit of Chinese tupelo is a dark blue oblong drupe that is readily eaten by birds. Taxonomy references place Nyssa either in Cornaceae (the dogwood family) or in its own family, Nyssaceae. The genus name Nyssa comes from Greek mythology and refers to a water (or rain) nymph named Nyssa (or Nysa), one of the nymphs who cared for Dionysus, god of wine, as a child (the location where the water nymphs sheltered Dionysus and where he invented wine is known as Mount Nyssa). The reference to water is the important bit, since this alludes to the preference of all Nyssa species for moist soils. The type species for the genus is in fact another North American species, N. aquatica, commonly called water tupelo or swamp tupelo because it grows in very wet sites. Like its American relative, N. sylvatica, Chinese tupelo prefers evenly moist, acidic soil but also tolerates somewhat drier conditions. The Arboretum currently holds just one specimen of Chinese tupelo (N. sinensis, accession 374-81-B), which grows near several other Nyssa accessions near Rehder Pond. This accession was received as seeds from China's Hangzhou Botanical Garden in 1981, but the exact provenance of their collection is unknown. Chinese tupelo has a fairly large range in central to southern China and an individual tree's cold hardiness may vary with provenance, but a probable hardiness rating would be through USDA Zone 6b (average annual minimum temperature 0 to -5 "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23446","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd1708128.jpg","title":"2015-73-2","volume":73,"issue_number":2,"year":2015,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Charles Edward Faxon: Botanical Draftsman","article_sequence":1,"start_page":2,"end_page":10,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25589","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160bb28.jpg","volume":73,"issue_number":1,"year":2015,"series":null,"season":null,"authors":"Pearson, Lisa; Dosmann, Michael S.","article_content":"Charles Edward Faxon: Botanical Draftsman Michael S. Dosmann and Lisa E. Pearson ear the Arnold Arboretum's Bradley Rosaceous Collection, at the south end of Meadow Road, lie three ponds whose names commemorate three staff members from the early years of the Arboretum: Jackson Dawson (propagator and superintendent), Alfred Rehder (taxonomist), and Charles Faxon (assistant director and botanical illustrator). These men, along with founding director Charles Sprague Sargent and explorer-botanist Ernest Henry Wilson, played central roles in shaping the Arboretum into the renowned institution that it remains today. Faxon's mark--in indelible ink no less--is the one we celebrate here. Charles Edward Faxon was born in the Jamaica Plain section of Boston, Massachusetts, in 1846, not far from the land that was to become the Arnold Arboretum in 1872. As a child, he developed dual interests in natural history and art. Much of his schooling in natural history was provided by his older brother Edwin Faxon (1823 Charles Edward Faxon JON HETMAN 3 assistant director at the Arboretum. In this position he was to curate the herbarium and organize the library, both of which were growing as quickly as the living collections. However, Faxon's primary charge was to assist Sargent with the Silva of North America by producing its illustrations. This seminal treatment, written by Sargent, spanned 14 volumes published between 1891 and 1902, and covered the known woody plants of the United States and Canada. Sargent-- the then \"dean of American dendrology\"--wrote eloquently and assertively about the various ligneous species, while Faxon brought the plants to life with painstaking detail and beauty in pen-andink. By the end of the project, some 744 plates for the Silva had been produced from Faxon's ink drawings. One fine example is his illustration of the vine maple (Acer circinatum), native to the Pacific Northwest. Faxon captured the full array of diagnostic characteristics necessary for identification, without whimsy, yet with an astonishing delicacy and A contemporary view of Faxon Pond (see inside front cover for a lantern slide from grace. In the forefront, the nearly the same viewpoint, circa 1920). eyes are drawn to a rounded wonderfully complementing Eaton's erudite leaf, the margins and primary veins boldly and text. Charles was responsible for a number of prominently outlined, as are the striking fruits the plates in Volume One and all of the plates from the same plane. The remaining leaf of this in Volume Two. branch, and those shown on the flowering and sterile branches in the background, are drawn in FAXON AT THE ARBORETUM lighter weights. When coupled to his subtle use of shading, the variable line weights effectively In 1879, Faxon became a botany instructor at create a depth of field, a sense of realism that Harvard's Bussey Institution, a school adjacent does not detract from the scientific purpose. to the Arboretum that was dedicated to the Magnified details of individual flowers, both agricultural and natural sciences. In 1882, C. S. Sargent hired him on a part-time basis as an male and female, as well as fruits, accompany 4 Arnoldia 73\/1 Charles Edward Faxon 5 6 Arnoldia 73\/1 Charles Edward Faxon 7 Top: A Charles Faxon drawing of Sequoiadendron giganteum (then known as Sequoia gigante) foliage and cone from Sargent's Manual of the Trees of North America. Bottom: A Charles Faxon drawing of Malus tschonoskii (then known as Pyrus tschonoskii) from Sargent's Forest Flora of Japan. Two M. tschonoskii specimens (accession 3678-A and B) grown from seed that Sargent collected in Japan in 1892 still grow in the Arboretum collections. 8 Arnoldia 73\/1 10 Arnoldia 73\/1 "},{"has_event_date":0,"type":"arnoldia","title":"Book Review: The New Sylva: A Discourse of Forest and Orchard Trees for the Twenty- First Century","article_sequence":2,"start_page":11,"end_page":16,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25588","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160b76f.jpg","volume":73,"issue_number":1,"year":2015,"series":null,"season":null,"authors":"Andersen, Phyllis","article_content":"The New Sylva: A Discourse of Forest and Orchard Trees for the Twenty-First Century BOOK REVIEW Phyllis Andersen The New Sylva: A Discourse of Forest and Orchard Trees for the Twenty-First Century Gabriel Hemery and Sarah Simblet Bloomsbury, 2014. 390 pages. ISBN 978-1-4088-3544-9 I n 1664 the Royal Society in London published Sylva, or A Discourse of Forest Trees, and the Propagation of Timber in His Majestie's Dominions. The author, John Evelyn (1620 12 Arnoldia 73\/1 Book Review 13 BIODIVERSITY HERITAGE LIBRARY Illustration of white oak (Quercus alba) by Pierre-Joseph Redout 14 Arnoldia 73\/1 Book Review 15 SARAH SIMBLET, FROM THE NEW SYLVA, WITH PERMISSION OF THE PUBLISHER Sarah Simblet's detailed illustration of an English (or pedunculate) oak (Quercus robur) seedling from The New Sylva. 16 Arnoldia 73\/1 "},{"has_event_date":0,"type":"arnoldia","title":"Filing A Missing Rose Claim: Jackson Dawson and the Arnold Rose","article_sequence":3,"start_page":17,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25590","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160bb6d.jpg","volume":73,"issue_number":1,"year":2015,"series":null,"season":null,"authors":"Whitacre, Benjamin","article_content":"Filing A Missing Rose Claim: Jackson Dawson and the Arnold Rose Benjamin Whitacre I cannot suggest to the earnest rose lover any finer summer pilgrimage than one to the Arnold Arboretum. --J. Horace McFarland, editor and publisher of the American Rose Annual (1917a) undreds of new plant cultivars, complete with gorgeous catalogue pictures and euphoric blurbs, are introduced each year. Few will still be around in a hundred years, and unfortunately sometimes the most innovative and influential are the first to fade away. Jackson Thornton Dawson, whose hybrids revolutionized rose culture more than any since the first repeat-blooming China roses, remembered the quick rise and precipitous fall of his roses. Writing in Country Life in America in 1911, the Arnold Arboretum's first superintendent detailed his once successful climbing roses before closing with a comment on `Arnold', his breakthrough hybrid whose subsequent decline and near extinction remains one of the unresolved puzzles of modern roses. \"A rich crimson, darker even than `G 18 Arnoldia 73\/1 COURTESY OF STEVE NYMAN Jackson Dawson and the Arnold Rose 19 NANCY ROSE the \"affinities\" between species with an ambitious program of Rosa accessions. The mix of innovation and raw material paid off. By the mid 1880s, Dawson had done the impossible, twice. He wed the diploid Rosa multiflora to the tetraploid `G 20 Arnoldia 73\/1 ARCHIVES OF THE ARNOLD ARBORETUM Jackson Dawson and the Arnold Rose 21 Darlington corrected \"Hartford Botanic Gardens\" to \"The Arnold Arboretum\" in 1917 and Jules Gravereaux, the director of the Roseraie de L'Ha 22 Arnoldia 73\/1 Jackson Dawson and the Arnold Rose 23 ARNOLD ARBORETUM HERBARIUM This herbarium specimen of `Arnold' (accession 857-79) was collected in 1994 when the cultivar still resided in the Arboretum's Bradley Rosaceous Collection. 24 Arnoldia 73\/1 Jackson Dawson and the Arnold Rose 25 `Little Tot', before 1900. A Rosa multiflora hybrid also listed as `Little Dot.' Dawson explained that `Little Tot' bloomed so much he could never get enough wood to propagate. `Minnie Dawson', 1896. An attempt to create a more fully double Rosa multiflora flower by backcrossing `Dawson' with the species. Named for Dawson's wife. `Pauline Dawson', 1916. Named for Dawson's daughter-in-law, who helped manage the family's Eastern Nurseries. *`Royal Cluster', 1899. `Dawson' 26 Arnoldia 73\/1 Jackson Dawson and the Arnold Rose 27 A Love Story Coded in Rosa Species Jackson Dawson was the key player in the Arnold Arboretum's rose breeding program, but Ernest Henry Wilson, the Arboretum's renowned plant explorer, also has a fascinating connection to roses. Wilson has not always been portrayed as a passionate husband; rosarians Douglas Brenner and Stephen Scanniello sketched him as a Ulyssean adventurer under the spell of his English patron, the heiress Ellen Willmott, willing to leave his wife Helen and newborn daughter Muriel Primrose to travel to the other end of the world for years at a time (Brenner and Scanniello 2009). Letters from Arboretum director Charles Sprague Sargent to Willmott show that she had a unique influence over Wilson. When Sargent failed to persuade Wilson to explore China for the Arboretum, he wrote an uncharacteristically seductive letter to Willmott, hoping she would intervene. She did (Sargent 1906). Her power over Wilson consolidated the Arboretum's legacy. The three later collaborated on Willmott's monumental tome, The Genus Rosa. During his time in Asia, Wilson collected as many as 2,000 species of plants previously unknown to the west, including Rosa helenae, or \"Helen's Rose,\" and R. murielae, named for his wife and daughter respectively. He named three finds for Willmott, among them a rose, R. willmottiae. Of the three roses, Rosa helenae caught on with gardeners. Where other musk roses fainted, Helen's Rose endured; Wilson suggested that a new breed of cold hardy Noisette-style roses be bred from it (Wilson 1916b). Canadian hybridizers took him up successfully (Thomas 1994). In 1914, shortly after helping Willmott finish The Genus Rosa, Wilson returned to Asia, this time to Japan. Helen and Muriel Primrose accompanied him on this and future expeditions. In 1927, after the death of Sargent, Wilson became keeper of the Arnold Arboretum, remaining one of the rose's most vocal proponents. He and his wife died together in an automobile accident on October 15, 1930. Over the following 30 years, botanists noted a close alliance between Rosa helenae and R. rubus, another species Wilson collected (Thomas 1994). At Kew, which had received both R. helenae and R. rubus seeds from the Arboretum in a lot labeled \"seed No. 431\" (Sargent and Wilson 1913"},{"has_event_date":0,"type":"arnoldia","title":"Midsummer Flurries","article_sequence":4,"start_page":28,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25591","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d1608126.jpg","volume":73,"issue_number":1,"year":2015,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"Midsummer Flurries Nancy Rose I n the steamy greenness of the midsummer landscape, white flowers can add a cooling touch, sort of a visual equivalent to a scoop of vanilla ice cream. Several hydrangeas have just such a refreshing floral display, including oakleaf hydrangea (Hydrangea quercifolia) and its cultivars. This species is native to the southeastern United States, growing from North Carolina to northern Florida and west to Tennessee and Louisiana, but it also grows well in other regions. It is recommended for USDA Hardiness Zones 6 to 9 (average annual minimum temperature -10 to 0 "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23445","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170bb6e.jpg","title":"2015-73-1","volume":73,"issue_number":1,"year":2015,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The History of Minimum Temperatures at the Arnold Arboretum: Variation in Time and Space","article_sequence":1,"start_page":2,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25587","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160b36b.jpg","volume":72,"issue_number":4,"year":2015,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"The History of Minimum Temperatures at the Arnold Arboretum: Variation in Time and Space Michael S. Dosmann G iven the original charge to cultivate \"all the trees, shrubs, and herbaceous plants, either indigenous or exotic, which can be raised in the open air,\" it's not surprising that the Arnold Arboretum has long been interested in documenting local climate and weather, particularly as they relate to plant hardiness. Early publications such as Garden and Forest and Arnoldia's predecessor, the Bulletin of Popular Information, are replete with notes of what did and did not survive New England's climate. Arnoldia continues that theme with annual summaries of the previous year's weather (see page 12 in this issue), often with notes on plant performance. One of the most innovative projects linking plants and climate was Alfred Rehder's creation of the first Arnold Arboretum Hardiness Zone Map, which was published in the first edition of his Manual of Cultivated Trees and Shrubs Hardy in North America (Rehder 1927). On this map, Rehder divided the United States into eight different zones based on the average minimum temperature of the coldest month. Then, using information about what survived the winters in Boston and other regions, he assigned plants in his Manual to particular Arnold Arboretum zones of maximum hardiness. This novel application was further updated and improved by the Arnold Arboretum, and later inspired and gave rise to the hardiness zone map (see page 9) created and now perpetuated by the United States Department of Agriculture (USDA). (See Del Tredici 1990 for a broader review, as well as Dosmann and Aiello 2013 for a brief discussion on the 2012 version of the map and its application to plant acquisition and collections planning.) It is important to bear in mind that the zone parameters in the Arnold Arboretum scheme were different from those in the USDA's, thus giving rise to confusion about a species' cold tolerance, particularly when a species was simply said to be \"hardy to Zone 6\" without further clarification--was it the Arnold's Zone 6 (average annual minimum temperature -5 to 5 Arboretum Microclimates 3 JORDAN WOOD Location of Weather Stations at the Arnold Arboretum of Harvard University 0 0.25 Mi Hunnewell Visitor Center N Dana Greenhouses Bradley Rosaceous Collection Bussey Hill 198 ft. Arboretum and Bussey Institution Greenhouses Explorers Garden Weld Hill Research Building Hemlock Hill 170 ft. Weld Hill 175 ft. Peters Hill 240 ft. Weather Stations Years Active Current 2009 4 Arnoldia 72\/4 Arboretum Microclimates 5 nate those plants unable to survive at those temperatures. It pays to be conservative when playing the hardiness game. Location, Location, Location In examining nearly a century of annual variation in minimum temperature at the Arboretum, one must bear in mind that those data were obtained from three separate and distinct locations, each with its own elevation and proximity to buffering buildings or canopies, as well as differences in aspect. And although we know that the present Weld Hill and Dana Greenhouse stations are sufficiently far away from buildings not to be influenced by them, we are not exactly sure where Judd's station was--it may have been somewhat protected. The Arboretum landscape comprises some 281 acres, with elevations that range from 44 feet (13.4 meters) above sea level in the Meadow by the Hunnewell Building to 240 feet (73.2 meters) on the summit of Peters Hill. Peters, Hemlock, and Bussey Hills each have their own character and microclimates distinct from surrounding areas. William Judd recorded daily weather notes, including maximum and minimum temperatures, from 1918 through 1946. The entry for February 9, 1934, (about half way down on the right) shows an extremely cold reading of -18 6 Arnoldia 72\/4 ARCHIVES OF THE ARNOLD ARBORETUM Arboretum Microclimates 7 MICHAEL S. DOSMANN cooling, warm air is lost to the atmosphere and cools. This forms a temperature inversion, where the denser, colder air then settles into the exposed areas at the lower elevations, creating what are often referred to as frost pockets. On this evening, the average minimum temperature of Raup's seven stations was -18.6 8 Arnoldia 72\/4 Arboretum Microclimates 9 of bowls--the frost pockets--had cold, dense air. Yet in other years, such as 2011 10 Arnoldia 72\/4 Arboretum Microclimates 11 ARCHIVES OF THE ARNOLD ARBORETUM KYLE PORT The weather station at the Dana Greenhouses, photographed by Arboretum plant propagator Al Fordham in the summer of 1969 (left), and the current state-of-the-art weather station located at the Weld Hill Research Building (right). for data collection. This season we are experimenting with a new set of loggers. The originals, while excellent in some respects, were nearing the end of their lifespan and required too much additional care; we have retired them. As part of her research, Arboretum Putnam Fellow Ailene Ettinger deployed a new set of loggers across an even wider swath of the landscape. These pendent loggers (Hobo 8KUA-002-08) are less intrusive in the landscape, easier to access and maintain, and are collecting temperature data at similar intervals. As I hope this article has demonstrated, a single landscape like the Arboretum's is marvelously variable. The year-to-year variation in elements such as temperature can be quite significant, particularly when compared across the Arboretum's unique nooks and crannies. I not only find this fascinating as a scientist, but as a dedicated horticulturist I am excited that ongoing data collection and analysis will allow us to best match the plants curated in the Arboretum with their optimum locations. Acknowledgements The deployment and maintenance of the data loggers and subsequent data wrangling have been no small task, and I wish to call attention to the many who have assisted in the venture over the years. Erik Youngerman, Sue Pfeiffer, Abby Hird, Jonathan Damery, Sam Schmerler, Stephanie Stuber, Joyce Chery, and Kyle Port did a lot of the heavy lifting out in the landscape with the loggers and data. Jordan Wood created the wonderful map integrating the old and new stations. Will Buchanan spent countless hours in the library putting the Raup and Judd data into spreadsheet form. Lastly, Mike Iacono not only provided data from the Blue Hill Observatory but also gave valuable comments on an earlier draft of this article. References Aiello, A. S. and M. S. Dosmann. 2007. The Quest for the Hardy Cedar-of-Lebanon. Arnoldia 65(1): 26"},{"has_event_date":0,"type":"arnoldia","title":"2014 Weather Summary","article_sequence":2,"start_page":12,"end_page":19,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25584","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160af28.jpg","volume":72,"issue_number":4,"year":2015,"series":null,"season":null,"authors":"Pfeiffer, Sue A.","article_content":"2014 Weather Summary Sue A. Pfeiffer KYLE PORT Horticultural Technologist Dennis Harris worked on clearing Meadow Road while intrepid visitors enjoyed the snowfall on February 5th. JANUARY started out cold and quite snowy. The first winter storm on the 2nd and 3rd delivered 15 inches of fluffy snow while the temperature dipped to -4 2014 Weather 13 peratures with highs in the 20s and lows in the teens and single digits. Seasonal temperatures returned on the 13th, and in the following week an additional foot of snow fell. Frigid temperatures returned and we finished out the month with lows in the single digits once again. The cold trend continued and the average high for the month fell four degrees below the historical average. MARCH brought a continuation of the cold pattern with lows in the single digits during the first week. The first signs of spring were evident on the 8th as temperatures warmed to the mid 50s, greatly melting the snow cover and reducing the icy, hard-crusted remnants of plowed snow. As the snow cover retreated, rabbit damage was visible on many shrubs. Precipitation during the month was scarce; we received two rainfall events on the 12th and 13th and on the 20th, amounting to less than an inch cumulatively. As temperatures warmed, the snow and ice continued to melt; by the 23rd, three days after the spring equinox, bare ground was visible as the snow cover had completely melted. A couple of cold and windy fronts moved in on the 22nd and 26th, both with average wind speeds of 16 mph and gusts reaching 36 mph, making it feel even colder. A storm arrived on the 29th bringing two days of consistent rain followed by a combination of rain, sleet, and hail as the storm lingered into the 31st. Over three inches of rain fell, making up for the lack of precipitation earlier in the month. It felt as if spring was right around the corner as spring ephemerals popped up from the warming soil. Despite rising temperatures, the month as a whole turned out to be colder than the historical average; both the average high and average low temperatures were 5 14 Arnoldia 72\/4 KYLE PORT 2014 Weather 15 Ups and Downs Significant temperature fluctuations can affect many natural processes, and we saw several examples in May 2014. This was an explosive year for fire blight (Erwinia amylovora), a bacterial disease that affects apples (Malus), pears (Pyrus), and a number of other rose family genera including mountain ash (Sorbus), hawthorn (Crataegus), firethorn (Pyracantha), and flowering quince (Chaenomeles). The warmer temperatures experienced early in the month followed by cooler conditions with regular rain provided ideal conditions for fire blight bacteria to spread throughout the collection. We also witnessed a substantial fish kill in Dawson Pond (the largest of our three ponds) over the Memorial Day weekend. This common natural event, observed across the region, was attributed to low oxygen levels in the water caused by temperature fluctuations. JULY was characterized by heat, humidity, and torrential downpours. We experienced several consecutive days of hot and humid weather, with the hottest day of the year (94 16 Arnoldia 72\/4 2014 Weather 17 of rain, officially making it the driest September since climate records began. These drought conditions, combined with the low rainfall amounts during late August, translated to accumulations of only 0.83 inches over a six-week period. The effects of this drought were apparent throughout the landscape; soils were extremely dry and the air was very dusty. Two of the Arboretum ponds almost dried up completely. Most A garter snake (Thamnophis sirtalis), the state reptile of Massachusetts, navigated through fallen leaves in mid-October. plants showed some signs of drought stress and severe stress was obvious on many plants. Leaves were flagging, some turning brown; many plants had already formed their winter buds and appeared to go dormant early. Because of these continued dry conditions, fall planting was postponed until the following spring. Despite all this, fall leaf color on maples (Acer), cork trees (Phellodendron), and birches (Betula) was exceptional throughout the landscape. OCTOBER was a warm and wet month. We started out with some much-needed precipitation from a storm that passed through on the 1st and 2nd, delivering well over an inch of rain. Sunny skies were prevalent as temperatures remained above average. A warm front moved through mid-month and we hit a high in the 80s on the 15th before temperatures returned to seasonal averages. The first nor'easter of the season arrived on the morning of the 22nd, bringing with it a welcome rain. As the storm intensified overnight, torrential downpours and high winds prevailed; recorded gusts peaked at 41 mph. A microburst (a small but intense downdraft of air) in the Centre Street Gate vicinity resulted in the complete loss of two accessions--a black hickory (Carya texana, accession 12892-A) along with a centenarian pin oak (Quercus palustris, accession 22896-E) were completely uprooted and broken below the base. Rain continued to fall until the 24th, delivering over three inches of precipitation. Other than the loss of the two large trees and damage to several nearby plants, the impact on the collection was minor with just some smaller branches down. Rain accumulation for the month was more than double that of the months of August and September combined! NOVEMBER began with a nor'easter on the 1st and 2nd; wind gusts reached 35 mph and we recorded an additional 3\/4 of an inch of precipitation equivalence which included a few hours of snow flurries on the 2nd. The snow created a beautiful juxtaposition in the landscape, but this did not last long as temperatures quickly warmed into the 60s. The growing season came to an end on November 10th when the first frost was recorded, ending the growing season at 202 days, the KYLE PORT Arnold Arboretum Weather Station Data 2014 Weather 19 SUE A. PFEIFFER Heavy rain in early December resulted in mulch washouts, especially on slopes like this one in the Lilac Collection. longest we have seen in over 7 years. Mid-November saw overnight temperatures dip well below freezing; this combined with rain and wind resulted in many trees dropping the remainder of their leaves, bringing an end to fall color. Another significant rainfall was recorded on the 17th, bringing over an inch and a half of rain. The last week of the month was very moist; we received 3 rain\/snow events accounting for almost 2 inches of precipitation equivalence. Overall, November was a wet and cool month; average temperatures were 2 degrees below normal and accumulated precipitation exceeded five inches. DECEMBER was a very wet and warm month; temperatures were 4 degrees above average and rainfall was abundant for the third month in a row. High temperatures during the first week fluctuated between the mid 30s and lower 60s with three storms depositing a total of almost two inches of rain. The next storm hit on the 9th, bringing wind gusts of over 40 mph and sustained winds at 18 mph--the highest recorded for the year. An additional three inches of rain fell, bringing the 10-day total to more than 5 inches. All of this rain left eroded gullies in gravel pathways and mulch washouts from planting beds, especially those in the lilacs. The rain subsided temporarily and temperatures remained above seasonal averages, allowing the grounds crew to accomplish much pruning and mulching. We recorded four additional rain events before we hit a high in the 60s on the 25th. These temperatures would not last as we ended the year with highs just below freezing. Little did we know what lay in store for the rest of the winter as we moved into 2015. Sue A. Pfeiffer is an Arboretum Horticulturist at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Hamamelidaceae, Part 2: Exploring the Witch-hazel Relatives of the Arnold Arboretum","article_sequence":3,"start_page":20,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25585","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160af6d.jpg","volume":72,"issue_number":4,"year":2015,"series":null,"season":null,"authors":"Gapinski, Andrew","article_content":"Hamamelidaceae, Part 2: Exploring the Witch-hazel Relatives of the Arnold Arboretum Andrew Gapinski I n \"Hamamelidaceae, Part 1\" we looked at just one genus, Hamamelis, in the witchhazel family. In Part 2, we'll study the other representatives of the family that are present in the Arnold Arboretum. It's helpful to start by looking at the evolutionary relationships among the genera in Hamamelidaceae. As mentioned in Part 1, witch-hazel (Hamamelis) displays showy flowers, each with four straplike petals. Several other genera within the family also have four-petaled flowers but they are found in warmer regions of the world and are not represented in the Arboretum, except for a lone specimen of Loropetalum (see page 26). In the past, these four-petaled genera were thought to be closely related on the family tree but recent DNA work is proving otherwise (Li and Bogle 2001). For example, when looking at the very similar appearing flowers of Hamamelis and Loropetalum it's easy to think they must be closely related (at one time both were included in the same genus), but in fact they are distant relatives found on separate branches of the family tree. The closest relatives of Hamamelis actually include genera such as Fothergilla, Parrotiopsis, and Parrotia (Li and Bogle 2001). Furthermore, the more advanced genera on each branch of the tree are those that have lost their showy, insect-attracting petals altogether, which is seen as an evolutionary shift from insect to wind pollination (Figure 1) (Li and Del Tredici 2008; Li et al. 1999). Among these aforementioned genera, Hamamelis is the oldest in evolutionary terms and is insect pollinated; Fothergilla and Parrotiopsis appear to represent an intermediate state in the transitional period and likely have both insect and wind pollination; and Parrotia, the most advanced, relies mainly on wind for pollination. Similar transitions take place on the other branches of the tree as well. We pick up here with the historical, taxonomic, and horticultural stories of the rest of the witch-hazel family starting with the closest relatives of Hamamelis. ANDREW GAPINSKI Fothergilla Fothergilla Gardeni[i] was introduced into English gardens one hundred and thirty years ago [1765], and judging by the number of figures that were published of it in Europe toward the end of the last and at the beginning of the present century, it must at that time have been a wellknown and favorite inhabitant of gardens from which it has now almost entirely disappeared, in spite of the fact that few shrubs present a more curious and beautiful effect than Fothergilla when it is covered with flowers. Its habit is excellent, too, and its foliage is abundant and rich in color. C. S. Sargent, Garden and Forest, 1895 The flowers of Loropetalum look very similar to those of Hamamelis, but the two genera are not closely related within the witch-hazel family. This is a flower of L. chinense on the Arboretum's sole specimen (see page 26). Hamamelidaceae, Part 2 22 Arnoldia 72\/4 NANCY ROSE 24 Arnoldia 72\/4 Hamamelidaceae, Part 2 MISSOURI BOTANICAL GARDEN, WWW.BOTANICUS.ORG 26 Arnoldia 72\/4 KYLE PORT 28 Arnoldia 72\/4 ANDREW GAPINSKI 30 Arnoldia 72\/4 32 Arnoldia 72\/4 HARVARD UNIVERSITY HERBARIA 34 Arnoldia 72\/4 "},{"has_event_date":0,"type":"arnoldia","title":"Pterostyrax hispidus, the Fragrant Epaulette Tree","article_sequence":4,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25586","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160b326.jpg","volume":72,"issue_number":4,"year":2015,"series":null,"season":null,"authors":"Thompson, Pamela J.","article_content":"Pterostyrax hispidus, the Fragrant Epaulette Tree Pamela J. Thompson I began my relationship with fragrant epaulette tree (Pterostyrax hispidus) when longtime Arboretum supporter and volunteer Elise Sigel brought me a lanky, homely specimen, wondering if I could give it a home. Elise couldn't recall its full botanical name (some sort of styrax?), and I failed to record even this. I planted it in my Milton garden, not knowing what I had, or how it might grow. Though I don't widely recommend this blind-faith landscape design strategy, in this case I've been delighted with the results. Pterostyrax hispidus is a deciduous tree native to Japan, specifically in the forested mountains of Honshu, Shikoku, and Kyushu. A member of the storax family (Styracaceae), it is closely related to the silverbells (Halesia). Though it can grow almost as broad as tall, reaching up to 50 feet (15.2 meters) in height and 40 feet (12.2 meters) in width as a tree, it is more often noted as a large multi-stemmed shrub reaching about 25 feet tall. In fact, it was the shrub form that Arboretum Director C. S. Sargent first saw in 1892 growing \"... wild in Japan on the banks of a stream among the mountains above Fukushima.\" The leaves of fragrant epaulette tree are oblong with a tapered point and have finelytoothed margins. They range from 3 to 7 inches (7.6 to 17.8 centimeters) long and 2 to 4 inches (5.1 to 10.2 centimeters) wide. Handsomely bright green above and gray-green below in spring and summer, the leaves turn yellowgreen to yellow in autumn before dropping. A truly remarkable feature of this plant is its profusion of 7- to 9-inch-long panicles of fringed, downward facing, white flowers that appear in mid to late June (in the Boston area). Hanging below the leaves, the flower clusters sway in the breeze, attracting multitudes of pollinators and giving off a delicate sweet scent. The inflorescences, reminiscent of the fringed epaulettes that once adorned the shoulders of military uniforms as a show of rank, give fragrant epaulette tree its common name. Through the summer, long clusters of indehiscent, bristly dry drupes develop, adorning the tree like bronze-chartreuse ornaments. These are most evident once the leaves have dropped, looking somewhat reminiscent of dangling sections of a DNA helix. The Arboretum's accession records for Pterostyrax hispidus reveal a history of human interest and persistence in growing this plant. The Arboretum acquired its first accession in 1880 from J. Veitch and Son in England. Over the next 130 years, the Arboretum acquired plants and seeds, including the 1892 accession collected by Sargent in Japan. Many of these acquisitions, though, were of garden origin or uncertain provenance. The Arboretum currently has 3 accessions (9 total plants) of Pterostyrax hispidus. Accession 218-60 came to the Arboretum as seed from the University of British Columbia, Canada, but with uncertain provenance. Accession 241-2008, received from Chiba University in Japan as seed, was wild collected in 2006 in Gunma Prefecture, Kanto District, about 20 miles northwest of Tokyo. The third accession, 843-76, came from the Academy of Sciences, Vacratot, Hungary, in 1976 and is also of uncertain provenance. Though it received the Royal Horticultural Society's Award of Garden Merit in 1993, Pterostyrax hispidus remains uncommon in the nursery trade. It is often listed as hardy to USDA Zone 4 (average annual minimum temperature -20 to -30"},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23444","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170bb28.jpg","title":"2015-72-4","volume":72,"issue_number":4,"year":2015,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Bark: From Abstract Art to Aspirin","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25581","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160a726.jpg","volume":72,"issue_number":3,"year":2015,"series":null,"season":null,"authors":"Begley, Eva","article_content":"ALL IMAGES BY PAUL AND EVA BEGLEY EXCEPT WHERE NOTED Bark: From Abstract Art to Aspirin Eva Begley T In the fading light of dusk, satiny bark curls on a greenleaf manzanita (Arctostaphylos patula) take on a purplish sheen. The bark of whitebark pine (Pinus albicaulis) is much finertextured than that of most pines and resembles an extreme close-up of an impressionist painting. o many people, bark is just the gray or brown stuff that covers tree trunks, but it's actually much more interesting than that. Woody dicotyledons and gymnosperms depend on their bark to keep insects and pathogens out. Bark also minimizes evaporation of water from trunks and branches. The fireresistant bark of giant sequoia (Sequoiadendron giganteum) grows up to 18 inches [45.7 centimeters] thick and has allowed some individuals to thrive for more than 3,000 years. Cork oak (Quercus suber), native to southwestern Europe and northwestern Africa, can also survive forest fires thanks to its thick bark. While functional for the tree, bark can be aesthetically pleasing for us. The bark of some trees shows surprising colors, including green, blue, and orange. It can be rough or smooth, stringy or flaky; it can peel away in long shreds or curl like chocolate shavings on an elaborate gateau. The textures and patterns in bark may remind you of abstract painting or sculpture, jigsaw puzzle pieces, or an old cable-knit sweater. Bark's charms are sometimes accentuated when festooned with lichens or providing a foothold for epiphytes. Anatomy of a Tree As a tree grows taller and adds more leaves and branches, its weight increases. To support the added weight, the trunk and branches grow in diameter. They do that thanks to a sleeve of almost-forever-young cells called the vascular cambium. During the growing season, these cells divide many times, mainly in a plane parallel to the surface of the trunk or branch. Cells produced on the inner side of the vascular cambium become xylem, which, as so-called sapwood, conducts water and minerals absorbed by the roots to the rest of the tree, then turns into the strong woody core of the tree--the heartwood, which is usually darker in color The trunks of giant sequoias (Sequoiadendron giganteum) are protected by thick layers of fibrous, fire-resistant bark. Bark 3 Mixture of dead cork cells and older, dead phloem Oldest, dead cork cambium Heartwood Sapwood Vascular cambium Xylem Older, dead cork cambium New, live cork cambium Phloem Bark What Is Bark? Botanists usually use the term \"bark\" to refer to everything outside the vascular cambium: phloem; phloem fibers; the innermost, live cork cambium and all its inner and outer derivatives; and older, dead cork cambia along with whatever else has accumulated outside the live cork cambium. The cork cambium and its products (that is, phellem and phelloderm) are collectively referred to as \"periderm.\" The live, deeper-seated components of the bark are sometimes called \"inner bark.\" than the sapwood. Cells produced on the outer side of the vascular cambium become phloem, which conducts sugars and other carbon-based nutrients throughout the tree. In temperate climates, the xylem and phloem formed early in each growing season usually contain lots of relatively large cells; cells formed later in the growing season are smaller. As a result, the xylem and phloem are built up of concentric rings, each ring constituting one year's growth. Phloem rarely lasts more than a few years (more on that in a moment). Xylem, however, can last well beyond the life of the tree in the form of standing snags or downed wood, or as lumber in buildings and furniture. Similar processes take place in roots. Once in a while, to keep up with the increasing girth of the tree, the cells of the vascular cambium divide in a radial plane. The phloem and most other cells outside the vascular cambium, though, have matured and aren't able to keep dividing or enlarging--they get stretched to the breaking point. That triggers the development of a new layer of squat, dividing cells, the cork cambium or phellogen, usually near the stem's surface. Like the cells of the vascular cambium, those of the cork cambium divide mainly in a plane parallel to the surface. (Interestingly, the cork cambium isn't necessarily active at the same time as the vascular cambium--the cork cambium seems to function more on an as-needed basis, perhaps in response 4 Arnoldia 72\/3 Bark 5 ally, that first layer of cork also gets stretched excessively and starts to crack. In cork oak, occasional cell divisions in a radial plane allow the cork cambium to keep pace with the growth in girth, but more commonly the first-formed cork cambium dies and new cork cambium forms deeper in the trunk or branch, sometimes even in the outer, older part of the phloem. In some species, each new cork cambium forms a complete sleeve; other species produce many small, overlapping patches of cork cambium, a bit like curling shingles on an old roof. Often, these later cork cambia are initiated right underneath cracks in the tree's surface, like internal bandages, ensuring that no crack gets deep enough to damage the living interior of the tree. This process is repeated over and over throughout the life of the plant. Eventually, a complex structure is formed, with everything outside the innermost, most recently formed cork cambium either dead or dying. Bark Variations The texture of the bark depends largely on the shape and location of successive cork cambia and on the types of cells \"trapped\" between them. Chinese or lacebark elm (Ulmus parvifolia), for example, has many overlapping, irregularly shaped cork cambia fairly close to the surface. Trees with deeper-seated cork cambia have rougher, craggier bark, like northern red oak (Quercus rubra) and tulip tree (Liriodendron tulipifera). Layers of thin-walled cells, whether the inner derivatives of the cork cambium or part of the phloem, are structurally weak, so bark characterized by such layers is likely to flake or peel off easily. Phloem sometimes contains lots of long, skinny, thickwalled but pliable cells, called fibers; as old phloem gets incorporated into the bark, these fibers give it a stringy texture. In some pines, the outer derivatives of the cork cambium consist of alternating bands of suberized cork cells The bark of lacebark elm (Ulmus parvifolia) has a jigsaw-puzzle-like pattern. This Garry oak, also known as Oregon white oak (Quercus garrayana), has deeply creviced bark. 6 Arnoldia 72\/3 Bark 7 bark that is plain gray in color, albeit with various textures. But then there's the aptly named paperbark maple (Acer griseum) with peeling sheets of cinnamon colored bark, Father David's maple (A. davidii) with its characteristic vertical white squiggles on a bright green background, and coral bark maple (A. palmatum `Sango-kaku'), a Japanese maple that adds color to winter gardens with its brilliant red branches. Bark's appearance often changes with age, and it's common for the bark of twigs and young branches to differ from that of older limbs. An extreme example is European white birch, in which the rough, gray to almost black bark near the base of the trunk forms a stark contrast to the creamy white bark higher up. And in aspen (Populus tremuloides), wherever the trunk has been wounded, be it by fungal attack, natural NANCY ROSE abscission of the lower branches as the tree gets taller, a bear climbing the tree, or lonely sheepherders or bored teenagers carving their names into the tree, the bark becomes black and fissured, very different from the tree's normally smooth, pale bark. Bark Beneficiaries Thick bark has some obvious benefits to trees, but the cracks and fissures in that bark can also provide good habitat for other species. Especially on rough-barked trees, enough soil, organic debris, and moisture can collect to fill minute pockets in which lichens, mosses, and larger epiphytes such as ferns and orchids can get a toehold. Often, different species of lichens and mosses grow on the upper and lower surfaces of leaning tree trunks and large limbs. Younger branches of coral bark maple (Acer palmatum `Sangokaku') are bright red. Black bears have left permanent calling cards on the trunks of this quaking aspens (Populus tremuloides). 8 Arnoldia 72\/3 Bark 9 Sapsuckers drilled multiple rows of holes in this white alder (Alnus rhombifolia). Extensive sapsucker drilling may partially girdle trees, which can eventually lead to the tree's decline. Acorn woodpeckers constructed a granary in this valley oak (Quercus lobata). The tree is now dead, but the presence of a few remaining slabs of bark full of the distinctive holes indicates that the birds started their work while the tree was still alive or at least still had bark on it. from one or two thousand to tens of thousands of acorn-sized cubbies, and each year the birds drill many more holes to replace those lost as limbs break off and old trees fall. In fall, the birds harvest ripe acorns from the branches of nearby oak trees (they rarely collect acorns that have already fallen to the ground), pry off the caps, and hammer the acorns into the predrilled holes. The flat end of the acorn, which provides a better surface for hammering, is almost always on the outside. If the first hole is too large or too small, the bird will try other holes until it finds one that is just the right size for a snug fit. The acorns provide an important food source for the family throughout the winter and early spring. Contrary to earlier belief, it seems that the birds feed directly on the acorns, not just on the insect larvae that sometimes infest them. Some mammals feed directly on bark. Porcupines and snowshoe hares like conifer bark. Moose will eat bark in winter if nothing more to their liking is available. Beavers, on the other hand, love bark, especially aspen (which is abominably bitter to human taste buds), but also other Populus species, willows (Salix spp.), birch, red-osier dogwood (Cornus sericea), and other species. I've even seen conifers (specifically, lodgepole pine, Pinus contorta subsp. murrayana) felled by beavers. During the growing season, the animals eat the buds, leaves, and twigs of these plants as well as the bark. In winter, bark is their primary food. Since beavers can't climb trees to reach the goodies up in the canopy, their solution is to gnaw down the entire tree. They are amazingly efficient at this: I once watched a beaver scramble out of an Ozark river and up a steep bank to a young 10 Arnoldia 72\/3 Bark 11 COURTESY OF AMORIM AND APCOR (PORTUGUESE CORK ASSOCIATION) The bark of cork oak (Quercus suber) is carefully hand-harvested. The bark regrows and can be harvested again in about ten years. to waterproof their homes. In fact, so versatile is the bark of paper birch that it was used for everything from canoes to kitchen funnels; as Moerman puts it, \"Nearly any kitchen utensil common to the white man could be duplicated in birch bark by the Ojibwe.\" The homes and barns of North America's European settlers were often roofed with the bark of American chestnut (Castanea dentata). Some of those buildings might have been painted using brushes made by boiling basswood (Tilia americana) bark in lye, then pounding it to extract its hemp-like fibers, a technique the settlers learned from Native Americans who made rope, sewing thread, and woven bags from basswood bark. The settlers probably wore shoes made of leather processed with tannins extracted from hemlock or oak bark, and some of their clothes may have been dyed with quercitron, derived from the yellow-orange inner bark of the black oak (Quercus velutina). Alone or in combination with mordants or other dyes, quercitron can yield colors ranging from bright yellow to warm browns. It was used commercially until well into the twentieth century, when cheaper synthetic dyes were discovered. Human health has also benefitted from certain chemical compounds in bark. To limit being incessantly munched by herbivores and damaged by insects, some plants produce chemical defenses. Some of these defenses are simply metabolic by-products, such as the calcium oxalate crystals that render the bark of some pines unpalatable to browsers. Others, such as various alkaloids, tannins, and cyanogens (which give cherry bark its distinctive bitter almond scent and cough-suppressing properties), require greater metabolic input and their synthesis consumes nutrients, but they provide valuable protection to long-lived plants. It's these same compounds that make the bark of some species medically useful. 12 Arnoldia 72\/3 Bark 13 wine jars among its uses. It takes a cork oak tree 25 to 40 years to build up a layer of cork thick enough to harvest, but the first harvest consists of hard, crumbly material good only for bulletin boards and insulation. If the cork is removed carefully, a new phellogen develops in the phloem 25 to 35 days later. The tree resumes cork production and can be harvested again 9 or 10 years later. Not until the third harvest, however, is the cork of sufficient quality for wine stoppers. The trees typically live 250 to 350 years, so each tree can be harvested many times. The practice of harvesting bark in cork oak forests actually helps preserve this unique ecosystem from land development so many conservation organizations promote the use of natural cork. And even though oenological research suggests that it doesn't really make much difference whether wine is sealed with natural cork, synthetic stoppers, or screw caps, yanking a plastic stopper out of a bottle just doesn't provide the same sort of tactile pleasure that pulling a real cork does. So pull a real cork, pour a glass, and drink a toast to bark. Further Reading Bugalho, M. N., M. C. Caldeira, J. S. Pereira, J. Aronson, and J. G. Pausas. 2011. Mediterranean cork oak savannas require human use to sustain biodiversity and ecosystem services. Frontiers in Ecology and the Environment 9: 278"},{"has_event_date":0,"type":"arnoldia","title":"A Dream Come True","article_sequence":2,"start_page":14,"end_page":21,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25580","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160a36d.jpg","volume":72,"issue_number":3,"year":2015,"series":null,"season":null,"authors":"Ashton, Peter Shaw","article_content":"A Dream Come True Peter Ashton T he possibility of being appointed director of the Arnold Arboretum in 1978 had come as a considerable surprise, but I jumped at it. Ever since my first professional appointment in 1962 as forest botanist in the Sultan of Brunei's government, I had been sending plant specimens to the Arnold as one of the six leading botanical research institutions both within and outside the Far East that specialize in the flora of East Asia, tropical as well as temperate. I respected the Arnold's scientific reputation in large part because of former Arboretum director Elmer Drew Merrill's astonishing achievements on the flora of the Philippines and southern China. Arboretum notables Ernest Wilson and Alfred Rehder were also well known to me and, as a life-long gardener and amateur horticulturist, the Arboretum's unique design by Frederick Law Olmsted intrigued me. Mary, my wife, and I will never forget our first glimpse of the Arboretum. During my interview, I sensed unhappiness among staff; morale was low. Mary was asked why she would wish to leave Scotland and her sheep; \"Why on earth do you wish to come to this place?\" quizzed another. Even the housekeeper in the fine old guesthouse at the faculty club, where we were accommodated on the Harvard campus, expressed the same feelings, and the (somewhat mythical) view that the Boston area had a crime level unimaginable in Aberdeen. When I arrived, curation and the living collections policies bore the mark of the celebrated horticulturist Donald Wyman who had been at the Arboretum from his appointment by tropical systematic botanist and director Elmer Drew Merrill in 1935 until his retirement in 1970. Wyman's interest had been in ornamental horticulture, reflected in his book Wyman's Gardening Encyclopedia, still the most comprehensive text specifically designed for American gardeners. The Arboretum then, as now, continued to sustain the keen interest and support of many members of the Garden Club of America and the Federation of Garden Clubs, as well as the ornamental nursery industry. But I was skeptical that Harvard and its upper administration really understood its fundamental scientific importance, nor the importance of its potential role within the university. Indeed, only one director following Charles Sprague Sargent, Karl Sax, had used the living collections in his research. But research universities focus on endeavors that advance scientific theory. The Arboretum's global herbarium collection, and with it the systematic botanists, had been removed to Harvard campus in Cambridge in the 1950s on the recommendation of a review chaired by Professor Irving H. Bailey. That decision alone led to nearly a decade of litigation between the University and the Association of the Arnold Arboretum, Inc. Harvard's adjacent Bussey Institute for plant research finally closed near that time, its distinguished faculty, scholars and researchers having been relocated to Cambridge two decades earlier in the 1930s. The Arnold Arboretum had become a backwater for the University, indeed \"an orphan institution\" within the broad missions of the University to educate and discover. Among faculty, Carroll Wood was alone in running a course based on the collections by our time, though Peter Stevens also used them later. Around the time I assumed my position, the Jamaica Plain-West Roxbury neighborhoods had been experiencing long decline, and this, too, had impacted the Arboretum. Trash collection had become a major activity for grounds staff, kids periodically drove beat-up automobiles off the summit of Peters Hill, while two corpses were discovered in our first year, one head-first down a road drain. So, there was no shortage of challenges, but that gave the job particular interest! Once I accepted this challenging position, it became my goal to reinvigorate the research functions of the living collections of the Arbo- ALL IMAGES FROM THE ARCHIVES OF THE ARNOLD ARBORETUM Peter Ashton 15 Peter Ashton in the greenhouse, 1983. Given the pristine appearance of the Arboretum today, it's hard to believe that it was once plagued by litterbugs and vandals. The photo above shows a trash-strewn slope in the Conifer Collection in 1973. retum. Colleagues in Cambridge had to be convinced that a systematic collection of specimen trees could be a resource for cutting-edge research. But first the living collections themselves had to be reviewed, and a new curatorial policy defined and executed, before a convincing case could be made. Because Sargent, on advice from Asa Gray, one of the world's leading botanists in his time, had established a systematic collection of woody plants, carefully selected and documented, the key was to bring this founding vision back to the fore. As I soon discovered, the Arboretum could then assume a unique role among gardens in Boston that complemented Boston's other two great living botanical and horticultural gardens: Mount Auburn Cemetery, a horticultural landscape focused on trees; and the Garden in the Woods, a native wildflower garden. Together, these three wonderful botanical collections could together offer the public a diversity of plants unequaled anywhere else in the New World, and in very few other places elsewhere. I realized that our collective objective should be to complement, rather than compete. My first quest, therefore, was to see the original Olmsted road plan and planting scheme. As Sargent had intended, the collections were laid out in such a way that a visitor could observe the families of trees hardy in the climate of Roxbury \"without alighting from his carriage.\" On inquiry, I discovered that the Arboretum library did not have the plans, nor was it clear where they could be found! But the old Olmsted firm buildings and archives still existed at Fairsted in Brookline, thanks to the interest and commitment of the landscape architect Joe Hudack. Arboretum archivist Sheila Connor spent a fortnight searching for the original plans in a garage full of Olmsted's 16 Arnoldia 72\/3 Peter Ashton 17 ily focused on bringing an appreciation of natural landscapes to the general public in city parks, university campuses, and in his involvement with the growing conservation movement. Harking back to Capability Brown, he exploited the majestic spaces of the new continent including the growing cities, and achieved what was unachievable in crowded Europe. This accomplishment can still be admired and cherished in Boston's Emerald Necklace. Olmsted's Arboretum plans revealed how he seamlessly combined his philosophy of landscape design with the Peter Ashton in his office at the Arboretum, 1983. requirements of a systematic their governor-general of the East Indies. Modbotanical collection. Bearing in mind that trees eled after the king of Prussia's garden Sans Souci within genera and even families share much (\"carefree\"), Buitenzorg was set in Bogor, the architecture in common, groves of tree families, rather than species, can achieve a similar town that was built as the colonial administrative center on the island of Java. The gardens effect in the landscape. But cultivars selected were reorganized and landscaped under Stamfor outstanding color or shape must be used ford Raffles, founder of Singapore, who, in his with utmost discretion. twenties, governed the Dutch East Indies for the Thus it became clear that the OlmstedSargent design and planting plan not only proBritish who had expropriated them during the vided an optimal solution to the design of an Napoleonic wars. The gardens became a scientific establishment thereafter, while remaining arboretum whose purpose was both to provide a public park. For me, with a decade in Borneo at a representative systematic collection for systematic and comparative research, but it is a the start of my career, the plant explorations of historic landscape for designers and planners: a Sargent and Engelmann west of the Mississippi park within which the public can both recreate River recalled the great Johannes Teijsmann. and learn. I realized that such a project remained Thanks to his intrepid explorations of Borneo unique. The Royal Botanic Gardens, Kew, are and Sumatra in leech-gorged clogs, the Buitenzorg gardens (now the National Botanic Gardens a historic landscape, but their land is uncompromisingly flat, denying the curving sweep of of Indonesia) hold the world's greatest collection of tropical woody plants. From the outset Olmsted's contour-hugging roads at the Arnold. they too had been meticulously documented Neither did Kew start with a clear accession and curated. And they are beautiful to look at, plan. The aim at the Arnold, to introduce at though nothing compared to the Arboretum! least three provenances of each taxon, to record And they have had a research laboratory on their location of collection, and to ensure nomenclatural verification with an herbarium voucher, grounds for over a century (though they, too, is known to me in only one other great ninerecently had their herbarium moved to Jakarta teenth century botanical garden, Buitenzorg, by unthinking biological policy-makers). which was originally established by the Dutch My prime objective, of returning the Arboretum to the fold of great research institutions as an ornamental garden around the palace of 18 Arnoldia 72\/3 Peter Ashton 19 Peter Ashton (center, seated) at a meeting in front of the Hunnewell Building, 1982. faculty appointments, rests with the faculty themselves. The university's schools have their own faculty and policies. But the allied institutions, such as the Arnold Arboretum, are in a no-man's land in which responsibility for faculty and research appointments has changed from time to time. Those allied institutions that are recognized as essential assets for FAS academic departments were in the best position, for their appointment priorities coincide. But the director of the Arnold Arboretum, clarified by the lawsuit of the fifties, reported directly to the university's president. Derek Bok, president at that time, was Peter and Mary Ashton in 1988 20 Arnoldia 72\/3 Peter Ashton 21 Without researchers on the staff who wished to avail of a laboratory, I sought to attract the interest of faculty in the several plant science departments in the universities of the Boston region. Thanks to some beneficent friends of the Arboretum, funds had been promised for construction of a modest lab. But new laboratories are normally approved at Harvard only where there is a potential or existing faculty to attract to them, or where a group of existing faculty campaign for one. Unfortunately, my own research in tropical tree biology could hardly be said to avail of our temperate living collections. Had I depended on the living collections in Jamaica Plain and Roslindale, a case could have been made as a condition of my appointment. Instead, a conclusion was reached at a meeting of the OEB Visiting Committee in 1988 that the Arnold Arboretum should retain a separate existence from the department and therefore FAS, and that no strong case therefore existed for faculty appointments on its staff. Lawrence Bogorad, a past president of the American Association for the Advancement of Science, alone continued to support my viewpoint: It was clearly time for someone more suitably placed to take up the challenge. Eddy Sullivan, educator and at that time vice-mayor in the City of Boston's mayor Kevin White's government, who had become a staunch supporter in my negotiations with the city, quipped, \"You don't have to worry, Peter; if it all fails, you can always go home to Ireland\"! Seen in this setting, it was no surprise that my successor as Arboretum director, Bob Cook, was not initially optimistic about the prospects of my case to embed the university's research back into the Arboretum. Bob had come from directing Cornell Plantations, which enjoyed a successful research and pedagogic relationship with academic departments in one of the leading universities in both fundamental and applied agricultural research. In the expected way, he arrived with a new broom. It was not long, though, before he came to realize the importance, even if against all odds, for building a laboratory at the living collections if they were to stand any chance of returning to Harvard's academic fold. Freed of faculty influ- ence as he was by the Arboretum's detachment from FAS, it is to Bob's great credit that with dogged determination he gained the support of the president's representatives in the administration. Those were the times of skyrocketing endowment values, and Bob's ambition came to vastly exceed my wildest dreams. But he--and the endowment--paid a heavy price when the recession of 2008 arrived. But the new laboratory building was nearing completion; it was fortunately too late to go back. Bob Cook should be remembered as the director who successfully brought the Arnold Arboretum back to a position where it could valuably contribute to Harvard's research and pedagogic mission, and in which it could reignite a major program in fundamental tree research--but this is his story to tell. For the first time in almost a century, the magnificent new Weld Hill Research Building might serve as a magnet for a new director, who could be a leader in a field that would avail of both them and what is now again the outstanding research collection of living trees in the temperate world. And so it has befallen! In spite of severe budgetary constraints, current Arboretum director William (Ned) Friedman has brought the new laboratory building to life with graduate students, with new faculty and classes availing of the living collections, and is attracting researchers from other institutions. Most importantly, thanks to a new generation of faculty in OEB and changing understanding in the Harvard administration, Friedman has been able to gain the university's support for advancing the Arboretum's scholarly mission in spite of current financial constraints. And in the spirit of the original intent, the public programs have been enriched by enhancing public appreciation of science. Regular research seminars have returned to the Arboretum, while the Director's Lecture Series is introducing increasing audiences to a variety of issues in the social as well as biological sciences. My dream has indeed come true, and with a flourish! Peter Ashton is Harvard University Bullard Professor Emeritus and was Director of the Arnold Arboretum from 1978 to 1987. He and Mary live in Somerset, England. "},{"has_event_date":0,"type":"arnoldia","title":"Lighting the Night: The Use of Pitch Pine and Bayberry in Colonial New England","article_sequence":3,"start_page":22,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25583","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160ab6f.jpg","volume":72,"issue_number":3,"year":2015,"series":null,"season":null,"authors":"Connor, Sheila","article_content":"Lighting the Night: The Use of Pitch Pine and Bayberry in Colonial New England Sheila Connor I TORCHES OF PINE n dark, small-windowed Colonial homes, the roaring fireplace brightened the room by day, and it often produced the only light available at night. Had domestic animals been abundant, the typical melted beef-suet or mutton-suet candles that the guildmakers produced in England would have been made. Tallow was scarce, however, and the inventive and resourceful settlers turned to materials ranging from extremely combustible meadow rushes soaked in lard to fish oil burned in shallow, wrought-iron holders, called Betty lamps, to illuminate their homes. These lamps An illustration of pitch pine (Pinus rigida) showing the cones still tightly closed, from A Description of the Genus Pinus by Aylmer Bourke Lambert, 1832. Pitch Pine and Bayberry 23 sputtered, smoked, and smelled unpleasant. A new method of lighting discovered by the colonists consisted of burning the resin-rich wood of a conifer that grew on the sandy coastal plains and ridges and in the sand barrens of river valleys. Pinus rigida earned the names candlewood and torch pine from the Europeans after they had observed how easily the Indians produced a bright flame by igniting several slivers of wood cut from its \"fat\" heartwood. The colonists referred to these sputtering torches that dripped pitch as \"splint lights.\" Whether growing in sterile seaside sands, where they are frequently bathed by salt spray, or rooted on exposed, windswept rocky hill tops, the torch or pitch pine thrives under adverse conditions. Easily blown over when young, a pitch pine eventually develops a root system that is substantial and deep enough to anchor it and to allow the tree to grow on an extremely dry site. Trees not more than four inches in diameter can have roots that penetrate to a depth of more than nine feet. Forest fires in these dry, windy habitats are devastating; however, not only do pitch pines survive, they often come to dominate the landscape after a fire. In New England, only Pinus rigida and the rarer P. banksiana, the jack pine--a tree of the Boreal Forest--are members of a group of conifers known as fire pines. These trees can withstand fire because they have evolved several specialized characteristics. All fire pines are pioneer trees--trees able to tolerate growing in full sun. Some have a high percentage of cones that remain closed until heat generated by fire melts the resin that glues the tips of their scales together, thereby releasing their seeds. These seeds remain viable inside the cone for many years, and they have the ability to germinate on soil totally lacking a humus component. The term \"serotinous,\" which means late-developing, describes the habit of bearing closed cones that contain viable seeds for many years. Jack pines retain their tightly closed cones for so long that they often become embedded in the wood of the tree's branches and can completely disappear as the branches thicken. Pitch pine's special adaptations include a thick, protective bark, some cones that remain closed, and the ability--unusual among conifers--to sprout A mature pitch pine cone that has opened and released its seeds. Cones may persist on the tree for years. from dormant buds on the main stem or at the base of the trunk if the tree is burned or cut. In New England, wherever the soil is exceptionally sandy, it is likely that pitch pines will be found. One of the few trees that can grow at the ocean's edge, flourish in salt marshes, and inhabit slowly moving sand dunes, Pinus rigida abounds on Cape Cod. Stunted oaks (black, red, scarlet, and white), along with the smaller post oak (Quercus stellata) and the Cape's ubiquitous scrub oak (Q. ilicifolia), are the common deciduous trees, but rising slightly above their crowns are the branches of the pitch pine, the true indicator of this sand-plain community. Usually reaching heights of less than fifty feet under the best of growing conditions, at thirty feet these pitch pines overtop the Cape's stunted forest canopy or form pure stands of low pine woods. Whether described as being New England `s most grotesque or most picturesque pine, a stand of P. rigida growing on a sandy hillside evokes an image of an untamed landscape. Pitch pines seldom grow straight; they twist this way and that. Their bark is remarkably rough and scaly, its color a very dark reddish gray-brown. Sparse, irregularly TIM BOLAND 24 Arnoldia 72\/3 Pitch Pine and Bayberry 25 NANCY ROSE Pitch pines growing on Cape Cod. 26 Arnoldia 72\/3 TIM BOLAND Pitch Pine and Bayberry 27 berries earlier than September 10th was outlawed in Connecticut beginning in 1724. Berry gatherers apparently ignored this legislation, however; and illegally collected berries before the authorized date. As they picked, the women and children noticed that their hands grew smooth as they acquired a thin film of wax from the berries. Inventive housewives saved some of the berries that they collected and filled cloth bags with them in order to grease the bottoms of their heavy flatirons. For candlemaking, the twigs and other debris that came home in the berry pails were removed, and the cleaned berries were placed in large cauldrons, covered with water, and heated and simmered for hours. A greenish, oily liquid floated to the top and solidified as it cooled. Repeated several times, this part of the process included straining the liquid through cloth to remove any impurities. Finally, a clear, solid cake of olive green wax resulted. The blue green water that remained was put to good use: homemakers used it to dye their homespun cloth. Patience and a steady hand came next. Dipping a wick twenty-five times or more into the remelted wax made a thin, tapered candle. Allowing each layer of wax to harden before the candle was dipped again meant that this process could take at least half an hour. Dipping several wicks at once saved time; only the size of the pot governed the number of candles that could be produced. Revolving candle stands that enabled the woman to dip several wicks at once decreased the time required, and tinsmiths made metal molds into which the heated wax could be poured, which eliminated the laborious dipping process altogether. It is no wonder that these highly prized and brittle candles, the finest light source available, were carefully stored in long, narrow boxes specifically made for holding candles. Not only were bayberry candles a useful domestic product that was saved for use on special occasions, they also became articles of trade in the colonies, and they were probably the first objects manufactured by women to be exported from New England. The English held these candles in highest regard, and they even tried to grow bayberries themselves. The French also hoped to establish bayberry plantations. However, neither the French nor the English succeeded in bringing Morella pensylvanica into cultivation on a large enough scale to support a candlemaking industry. Sheila Connor is the former Horticultural Research Archivist at the Arnold Arboretum. This article is adapted from New England Natives by Sheila Connor, Harvard University Press, 1994. 36673667 U.S. POSTAL SERVICE STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION (Required by 39 U.S.C. 3685) 1. Publication Title: Arnoldia. 2. Publication No: 0004"},{"has_event_date":0,"type":"arnoldia","title":"Erable de Montpellier, the Montpellier Maple","article_sequence":4,"start_page":28,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25582","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160a76b.jpg","volume":72,"issue_number":3,"year":2015,"series":null,"season":null,"authors":"Urban-Mead, Katherine","article_content":"Erable de Montpellier, the Montpellier Maple Katherine Urban-Mead L ast year I declared I could never love any other tree as much as a sugar maple. After accepting a several-month ecology internship in Montpellier, France, I bid a teary adieu to the stunning October foliage around my Hudson Valley home. Then I stepped off the airplane into a new world of dusky gray and gnarled Mediterranean greens. Ancient olive trees stand like statues in the roundabouts; streets are dotted with palms, cypresses, and occasional figs; tightly-pruned planetrees line esplanades and bike paths alike. There is no maple syrup here. On my first day at work, I climbed a rickety external staircase to the third floor, and with some confusion saw samaras waving from an unfamiliar tree growing alongside the stairs. Paired samaras (one-seeded fruits with papery wings) are characteristic of the maples (Acer), a group of plants I had worked with as a horticultural intern at the Arboretum last year. During my internship I had puzzled over hawthorn maple (A. crataegifolium) and communed with paperbark maple (A. griseum), but had never taken time to get to know the species that I now greeted with great glee. It was not a sugar maple, but instead the aptly-named Montpellier maple, Acer monspessulanum. After my joy at finding a local maple subsided, I had to admit that the Montpellier maple is not a particularly elegant tree. It is sometimes referred to as a shrub (arbuste in French), with an average height of only 15 to 25 feet (4.6 to 7.6 meters). Its slow growth and small trunk, frequently branched into several stems, give it a craggy feel characteristic of many Mediterranean region trees. Montpellier maple's leathery three-lobed leaves are rounded and smoothedged, are borne on long petioles, and are only 1.5 to 2.75 inches (4 to 7 centimeters) wide and 1.25 to 2 inches (3 to 5 centimeters) long. By mid-November the morning chill in Montpellier had become crisper; the endearing leaves of the tree I pass each morning turned first yellow then red. Finally brown, they fell and were scattered through the halls by passing boots. In the spring, Montpellier maple bears small, bright greenish yellow flowers that open earlier than its leaves, followed by the parallel-winged samaras frequently tinted pink or red and maturing to tan. This drought-tolerant species handles occasional cold and persists in USDA hardiness zones 5 to 9 (average annual minimum temperatures -20 to 30 "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23443","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170b76e.jpg","title":"2015-72-3","volume":72,"issue_number":3,"year":2015,"series":null,"season":null},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23441","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170b36e.jpg","title":"2014-72-2","volume":72,"issue_number":2,"year":2014,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Hamamelidaceae, Part 1: Exploring the Witch-hazels of the Arnold Arboretum","article_sequence":1,"start_page":2,"end_page":17,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25577","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25e856b.jpg","volume":72,"issue_number":2,"year":2014,"series":null,"season":null,"authors":"Gapinski, Andrew","article_content":"Hamamelidaceae, Part 1: Exploring the Witch-hazels of the Arnold Arboretum Andrew Gapinski H KYLE PORT amamelidaceae, the witch-hazel family, includes approximately 30 genera representing around 100 species of deciduous trees and shrubs. Members of the family are found in both temperate and tropical regions of North and Central America, Eastern Asia, Africa, the Pacific Islands, and Australia. The Arnold Arboretum has a rich history with the family, from plant exploration to the naming and introduction of its members to cultivation. The Arboretum's Hamamelidaceae collection, which currently comprises ten temperate region genera, can be found in groupings throughout the Arboretum landscape. Specific locations Many witch-hazels display attractive fall color; seen here, Hamamelis Hamamelidaceae, Part 1 4 Arnoldia 72\/2 6 Arnoldia 72\/2 ARNOLD ARBORETUM 10 Arnoldia 72\/2 MICHAEL DOSMANN Hamamelidaceae, Part 1 ANDREW GAPINSKI 14 Arnoldia 72\/2 16 Arnoldia 72\/2 "},{"has_event_date":0,"type":"arnoldia","title":"Did American Chestnut Really Dominate the Eastern Forest?","article_sequence":2,"start_page":18,"end_page":32,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25576","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25e8526.jpg","volume":72,"issue_number":2,"year":2014,"series":null,"season":null,"authors":"Foster, David R.; Faison, Edward K.","article_content":"Did American Chestnut Really Dominate the Eastern Forest? Edward K. Faison and David R. Foster \"The American chestnut once comprised 25% or more of the Native Eastern Hardwood Forest.\" American Scientist (1988) \"Chestnut was perhaps the most widespread and abundant species in the Eastern United States since the last glaciation.\" USDA Forest Service Southern Research Station General Technical Report General Technical Report SRS-173 (2013) \"Before the turn of the century, the eastern half of the United States was dominated by the American chestnut.\" American Chestnut Research and Restoration Project, SUNY College of Environmental Science and Forestry (2013) A long with the bison and the passenger pigeon, the American chestnut forms an iconic triumvirate of the grandeur of the American wilderness and the devastation that human activity wrought upon it over the past three centuries. Just as the bison was the preeminent large mammal on the continent and the passenger pigeon the most abundant bird, so is chestnut often described as having dominated the eastern forest (or across its geographic range) prior to its destruction by an introduced Asian chestnut blight. By all accounts chestnut was a magnificent and invaluable tree. It was among the fastest growing, tallest, and widest-trunked trees in the eastern United States. The strength, straight grain, and decay resistance of its wood made it ideal for framing, finished lumber, and fencing, and its regular production of nuts provided abundant food for native and European peoples, domestic livestock, and diverse wildlife. But was it really the dominant tree in the eastern forest? Dominant species, in the words of forest ecologist E. Lucy Braun, are \"those trees of the canopy, or superior arboreal layer, which numerically predominate.\" Given American chestnut's purported prior dominance in the eastern deciduous forest, we would expect the tree to have ranged widely across the East relative to other common tree species and to occupy a superior place in written accounts by early naturalists and explorers, early land survey records, forest surveys of the early twentieth century, and the paleoecological record. In fact, these sources reveal a very different story. Accounts by Early Explorers and Naturalists Accounts by foresters about chestnut's abundance at the turn of the twentieth century have been widely cited in the scientific and popular literature as evidence of the tree's former dominance. Descriptions of chestnut by naturalists and explorers at the time of European settlement, on the other hand, are rarely cited. Early written records must be used with caution, given that they were often written by non-botanists and provide a potentially biased assessment of previous forest conditions (Whitney 1994). Nonetheless, these descriptions--particularly if they correspond with other available lines of evidence--provide valuable eyewitness accounts of eastern forests prior to their widespread modification by Euro- American Chestnut 19 FOREST HISTORY SOCIETY, DURHAM, NC DAVID R. FOSTER (Left) A large American chestnut photographed in the Monongahela National Forest, West Virginia, in 1923. (Right) Foliage of American chestnut (Castanea dentata). pean settlement. Below are selected quotations that reference chestnut and other species by some of the more important early explorers and naturalists in the Eastern United States. John Smith, New England coast (early 1600s): \"Oke [oak], is the chiefe wood, of which there is great difference in regard of the soil where it groweth; fir, pine, walnut, chestnut, birch, ash, elm ..., and many other sorts.\" (Smith 1616) Colonel William Byrd, Virginia (1737): \"chestnut trees grow very tall and thick, mostly, however, in mountainous regions and high land ...\" (Bolgiano and Novak 2007) William Bartram, northern Alabama A KILLER ARRIVES Chestnut blight (Cryphonectria parasitica) was first discovered in 1904 in a stand of American chestnuts (Castanea dentata) in New York's Bronx Zoological Park, perhaps arriving on imported nursery stock of Castanea crenata from Japan. Subsequent investigation determined that the blight arrived in the late nineteenth century, as evidence suggested that American chestnuts on Long Island had been infected as early as 1893. The effects of the blight were immediate and devastating, often killing mature trees in 2 to 3 years. By 1906, the blight was detected in New Jersey, Maryland, and Virginia and continued to spread rapidly, reaching Pennsylvania in 1908 and North Carolina by 1923. All government efforts to contain or eradicate the blight failed, and ceased entirely by 1915. By the early 1940s the destruction of the American chestnut throughout its 300,000-square-mile range was complete. The blight spreads by wind-borne fungal spores that invade the tree through cracks or injuries in the bark, killing the cambium and eventually girdling the tree. The roots generally survive the blight, however, and continue to produce sprouts that are eventually killed again before reaching reproductive age. In effect, the chestnut blight converted a once towering overstory tree into an understory shrub. An American chestnut in Connecticut succumbing to chestnut blight, from the image collection American Environmental Photographs, 1891 American Chestnut 21 A large white oak (Quercus alba) photographed near New Lenox, Illinois, from the image collection American Environmental Photographs, 1891 22 Arnoldia 72\/2 American Chestnut 23 CHESTNUT Pre-Colonial Relative Abundance Absent 0 to 2.5% 2.5 to 5% 5 to 10% 10 to 20% 20 to 40% > 40% BEECH Pre-Colonial Relative Abundance Absent 0 to 2.5% 2.5 to 5% 5 to 10% 10 to 20% 20 to 40% > 40% American chestnut abundance compared with American beech and eastern hemlock abundance in the Northeast at the time of European settlement as determined by early land survey data (Thompson et al. 2013) of trees in a single town. In contrast, beech comprised 22% of trees across the region; oaks, predominantly white oak, 17.5%; and hemlock 11%. Two decades ago, forest historian Gordon Whitney compiled maps of tree species abundance from land survey data across the midwestern United States. Data from about 100 counties or townships across eight states of the upper Midwest reveal that chestnut was never the dominant tree, comprising 5 to 15% of trees in a small section of Ohio and 0 to 4% of trees in the rest of the region. In contrast, beech and especially white oak were frequently the dominant tree, often comprising 25 to 65% of all trees. Limited early land survey data from the southern regions of the eastern forest also portray chestnut as a secondary species. Chest- HEMLOCK Pre-Colonial Relative Abundance Absent 0 to 2.5% 2.5 to 5% 5 to 10% 10 to 20% 20 to 40% > 40% 24 Arnoldia 72\/2 American Chestnut 25 A white oak (Quercus alba) in New Braintree, Massachusetts. nut was the first-ranked species in only one of 15 locations, whereas white oak was the firstranked tree in five of 15 locations (see Table on facing page). Early Twentieth Century Forest Surveys E. Lucy Braun conducted and compiled extensive forest surveys and observations across 120 counties of the eastern forest in the early twentieth century. Her data were predominantly gathered from \"original\" forests and thus fill in gaps in the witness tree studies, particularly in regions such as the Cumberland Mountains of Kentucky and the Blue Ridge Mountains of North Carolina and Tennessee. Although Braun acknowledged her unequal coverage of different regions, her work remains by far the most comprehensive assessment of the eastern deciduous forest, including American chestnut's abun- dance, at the time of the chestnut blight. Her surveys and data tables reveal that chestnut was a tree of surprisingly limited dominance. Chestnut was dominant (the most abundant canopy tree) in at least one survey in only 15 of the 120 counties (12.5%) sampled by Braun and others. Sugar maple, white oak, and hemlock were all dominant species in over 20% of the counties sampled, and beech was a dominant tree in over 40% of the counties sampled. In fact, Braun's data suggest that chestnut was not even the most abundant tree within its own geographic range: beech was a dominant species in at least one survey in almost half (48%) of the counties sampled in chestnut's range, whereas chestnut was a dominant tree in less than a quarter (23%) of the counties sampled. American chestnut was spectacularly abundant in some locations. On north slopes in Joyce JOHN S. BURK 26 Arnoldia 72\/2 American Chestnut 27 AMERICAN CHESTNUT WHITE OAK Chestnut Range White Oak Range Counties Sampled Dominant Present Absent Counties Sampled Dominant Present Absent American chestnut's geographic range and extent of dominance compared to that of white oak and American beech in the early twentieth century. Data compiled by Braun (1950). ability to sprout vigorously from cut stumps, including those of large diameter and advanced age, made it better adapted to intensive logging than any other hardwood tree including oaks. As the early Connecticut foresters Hawley and Hawes (1912) wrote, \"this sprouting capacity of the species is its strongest characteristic and the one by which with each successive cutting it gains in the struggle for existence with the rival inmates of the woodlot.\" Interestingly, chestnut's sprouting capacity was much more prominent in the Northeast than in the southern parts of chestnut's range. In heavily cutover forests of northern New Jersey and southern New England, chestnut increased from 5 to 15% of the forest during the early colonial period to an estimated 50% of the standing timber in Connecticut. Because Braun focused AMERICAN BEECH Beech Range Counties Sampled Dominant Present Absent 28 Arnoldia 72\/2 American Chestnut 29 on \"original\" forests in her surveys, she largely avoided surveying the cutover southern New England region so her data probably underestimate chestnut's abundance in the Northeast. But it's important to remember that southern New England represents a small fraction of chestnut's range and the eastern forest overall. SPATIAL SCALE Spatial scale refers to the size or extent of the area under consideration. A stand is a relatively small area of forest that is spatially continuous in structure and composition and is exposed to similar soil and climatic conditions. In paleoecology the size of the catch basin (e.g., lake, pond, swamp, or small hollow) determines the distance from which pollen in the sediments originates. Sediments from a small forest hollow will contain pollen from vegetation growing predominantly in the immediate stand (a \"stand scale\" investigation), whereas sediments from a large lake are dominated by pollen from the broader landscape up to 20 miles away. The Last to Arrive: Chestnut Since the Last Ice Age Fossil pollen records in the Eastern forest enable reconstruction of vegetation communities and tree species that have dominated forests over the past 15,000 to 50,000 years. In formerly glaciated areas such as the Northeast, pollen records provide a chronological record of recolonization of forest vegetation after glacial melt some 15,000 to 20,000 years BP (before present). In southern New England, ash (Fraxinus), birch (Betula), ironwood (both Ostrya and Carpinus, whose pollens are indistinguishable from each other), and oak arrived first, followed by maples; deciduous forests replaced coniferous forests about 9,000 years BP. Beech arrived about 8,000 years BP, and hickory about 6,000 years BP. Not until about 2,000 years BP does chestnut pollen appear in the sediment record, earning chestnut the distinction of being the last major tree species to recolonize the region DANILO D. FERNANDO, SUNY-ESF A micrograph of American chestnut pollen. after deglaciation (Davis 1983). When chestnut finally does appear in the sediment record, it generally doesn't exceed about 4 to 7% of the pollen types across the region with the exception of one record in northwestern Connecticut where it reaches 18 to 19% (Paillet 1991, Oswald et al. 2007). In contrast, oak pollen consistently comprises 40 to 60% of the pollen and beech 5 to 20%. Interestingly, chestnut does achieve great dominance (40 to 70%) at the stand scale in a few local New England pollen records (Foster et al. 1992, 2002), exemplifying the importance of spatial scale when considering the abundance of this species. What accounts for chestnut's late arrival to New England? One possible reason is that the climate of the Northeast throughout much of the Holocene was too dry for chestnut. Other researchers have posited a lack of favorable well-drained germination sites in southern New England after deglaciation, or too much lime in the soil that took millennia to leach away. Chestnut is also self-sterile unlike many other trees that are self-fertile, and thus the chances of establishing new populations were much lower for this tree. Whether dispersal or environmentally limited, it is clear that 30 Arnoldia 72\/2 American Chestnut 31 chestnut's great abundance (40 to 45%) in a few southern Appalachian pollen records analyzed by the Delcourts and stand-level records from Massachusetts are consistent with twentieth century forest surveys in which chestnut achieved great dominance in some landscapes and topographic positions, but generally not at broader scales. Abrams, M. D., D. M. McCay 1996. Vegetation-site relationships of witness trees (1780 32 Arnoldia 72\/2 "},{"has_event_date":0,"type":"arnoldia","title":"Reading Tree Roots for Clues: The Habits of Truffles and Other Ectomycorrhizal Cup Fungi","article_sequence":3,"start_page":33,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25578","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25e896f.jpg","volume":72,"issue_number":2,"year":2014,"series":null,"season":null,"authors":"Healy, Rosanne","article_content":"Reading Tree Roots for Clues: The Habits of Truffles and Other Ectomycorrhizal Cup Fungi Rosanne Healy H ere's something to ponder: The health and regeneration of grand old oaks (Quercus) and majestic pines (Pinus) is dependent on the well-being of tiny fungi that associate with the trees' roots. Such small organisms have a big role to play not only for oaks and pines but also for many other trees that rely on their fungal partners to get them through lean and dry times. An estimated 86% of plant species benefit from (or are even dependent on) fungal root associates that transfer water and nutrients to the plant in exchange for carbohydrates (Brundrett 2009). Carbohydrates from plants are the result of atmospheric CO2 (carbon dioxide) fixation through photosynthesis and subsequent processes, which the fungi are incapable of doing. The fungal root associates are the mycorrhizal (myco=fungus, rhiza=root) fungi. They can be roughly sorted into two types based on how they associate with the roots. One type is mostly invisible to us because their hyphae are inside the root (endomycorrhizae), and the other can be seen as a mantle surrounding the root tip (ectomycorrhizae). The endomycorrhizal fungi are root associates of the vast majority of herbaceous plants and certain tree species. This article focuses on ectomycorrhizal fungi, which grow mostly in association with trees rather than herbaceous plants. They make their presence known to us not only because we can see them on tree roots but also because we see their fruiting bodies, particularly from midsummer into fall here in New England. Trees such as the red oaks (Quercus rubra) and eastern white pines (Pinus strobus) seen here benefit from ectomycorrhizal fungi. ALL IMAGES BY THE AUTHOR EXCEPT WHERE NOTED The color and \"furry\" appearance of this ectomycorrhizal red oak root tip are from the fungal symbiont, a Scleroderma fungus. NANCY ROSE 34 Arnoldia 72\/2 Ectomycorrhizal Cup Fungi 35 RICHARD SCHULHOF Research indicates that the ample foot paths, mowed lawns, and sparse understory in the Arnold Arboretum will favor Pezizales fungi on the root tips of the ectomycorrhizal trees. Sporemats of truffle fungi Pachyphlodes sp. nov. (left) and Tuber sp. nov. (right). An example of a sporemat and the truffle (Pachyphlodes ligericus) that its fungal barcoding sequence matches. ing trees, native shrubs, vines, and herbs. The ground under the trees is covered by woody and leafy litter, and under that layer is a deep organic layer composed of roots, soil, and partially broken down organic matter that together form a dense mat that requires a knife to cut through it. Compared to the forest habitat, there is not much in the Arboretum habitat to obstruct the passive transfer of fungal spores produced on the soil surface to roots and mycelia in or below the organic layer. This is possibly an important feature for the cup fungi because in order to fruit, the hyphae of outcrossing species such as Tuber must come in contact with a compatible mating type nucleus in another hypha. This is in contrast to most ectomycorrhizal basidiomycete species that form their mycelia with both nuclei soon after germination of their sexual spores. How do compatible mating types of truffles get together if the mycelia are underground? Perhaps the sporemats on the soil surface play a role in this event. If so, mating may be facilitated in an environment such as that found in the Arb o retu m o v er th at found in a forest. 36 Arnoldia 72\/2 Ectomycorrhizal Cup Fungi 37 units (MOTUs) from Harvard Forest and 56 MOTUs from the Arboretum, 17 of which overlapped in both sites. Some MOTUs could be matched to sequences in GenBank from described species or at least sequenced fruit bodies. Russula species were the most frequently sequenced in both habitats with 32 MOTUs. A number of our other This Russula fungus (fruit body and root tip shown) has a sequence that matches root sequences matched Russula tips in this study, as well as root tips and fruit bodies from a 2006 study by Don Pfister and Sylvia Yang in which they determined that many Russula species are exploited by sequences from a previous the Indian pipe plant, Monotropa uniflora. study by Don Pfister and Nearly equal numbers of Ascomycete MOTUs Sylvia Yang, but not sequences of any described were sequenced from each site. However, there species. A distant second place for most commonly sequenced genus was Cortinarius (14 was little overlap in species. It is particularly MOTUs) followed by Lactarius (9 MOTUs). interesting that the Pezizales had significantly Even less common (genus followed by MOTUs greater species richness and number of root within parentheses): Amanita (4), Boletus (1), tips in the Arboretum (10 MOTUs) than in the Forest (3 MOTUs). The cup fungi detected on Byssocorticium (1), Clavulina (4), Craterellus roots in the Arboretum included Hydnotrya, (1), Entoloma (3), Inocybe (4), Laccaria (1), Piloderma (1), Pseudotomentella (1), Scleroderma four species of Pachyphlodes, three species of (2), Sistotrema (1), Strobilomyces (1), TomenTuber, and two root tip sequences that have no tella (7), Trechispora (1), and Tylopilus (1). match to a fruit body sequence. From Harvard Ectomycorrhizal ascomycete fruiting bodies (above) and their root tips (below) from (left to right) Elaphomyces muricatus, Pachyphlodes sp. nov., and Tuber separans. 38 Arnoldia 72\/2 Ectomycorrhizal Cup Fungi 39 The fruiting body and root tips of the newly named Tuber arnoldianum. again on the same tree, and is in the process of looking for it on other trees in the vicinity. A third interesting story involves another Tuber species. We detected a species (termed \"species 46\" by Tuberaceae expert Gregory Bonito, a mycologist at the Royal Botanic Gardens in Melbourne, Australia) on the roots of several trees scattered throughout the Arboretum, as well as from one of the trees sampled in Harvard Forest. Our sequences match those for an undescribed species, known previously only from orchid root tips in New York and red oak root tips from an urban area in New Jersey. We were fortunate to recover some fruiting bodies from the Arboretum so that we will now be able to describe this taxon. The Arnold Arboretum staff has chosen the name Tuber arnoldianum for this truffle. While data are still being gathered, enough has been analyzed at this point (985 root tip sequences from 24 trees in each site) that I expect the pattern of Basidiomycete to Pezizales MOTUs in the two sites to hold up. This pattern continues to support the hypothesis that Pezizales are more prevalent in managed woodland sites such as the Arboretum. We can't be certain of the determining factors for this pattern, but refining the experimental parameters will help to zero in on those factors that are correlative. The well documented history of each accessioned tree, the ease of access to the rich information regarding Arboretum vegetation, and the encouragement and support of research by the staff at the Arnold Arbore- tum and Harvard Forest make these sites ideal for helping to resolve some of the outstanding questions regarding the ecology of ectomycorrhizal cup fungi. References Brundrett, M. C. 2009. Mycorrhizal associations and other means of nutrition of vascular plants: understanding global diversity of host plants by resolving conflicting information and developing reliable means of diagnosis. Plant Soil 320: 37"},{"has_event_date":0,"type":"arnoldia","title":"The Castor Aralia, Kalopanax septemlobus","article_sequence":4,"start_page":40,"end_page":40,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25579","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d160a328.jpg","volume":72,"issue_number":2,"year":2014,"series":null,"season":null,"authors":"Port, Kyle","article_content":"The Castor Aralia, Kalopanax septemlobus Kyle Port K alopanax is a monotypic genus in Araliaceae, the ginseng family. The lone species, K. septemlobus, is a dominant tree in northeastern Asia (Japan, China, Korea, the Russian Far East) where it is valued for the ethnopharmacology of its plant parts and its timber quality. Across Korea, overuse has threatened some wild populations and there are now calls to protect the species. Castor aralia is a large deciduous tree that can grow to nearly 100 feet (about 30 meters) tall and has an average trunk diameter of about 40 inches (about 100 centimeters). Its stems are armed with stout prickles that yield to thick, deeply furrowed bark with age. It has very large (to 14 inches [36 centimeters] in diameter), longpetioled, 5- to 7-lobed leaves that may turn brilliant greenish yellow in autumn. Castor aralia bears large, wide (to 12 inches [31 centimeters] in diameter) inflorescences with numerous small umbels of white flowers that open in August and September here, providing late season nourishment to an assortment of pollinators. Successful pollination yields abundant blue-black fruits that are retained into winter. A single castor aralia plant was sent to the Arnold Arboretum in January 1881 by Alphonse Lavall "},{"has_event_date":0,"type":"arnoldia","title":"Seeing the Lianas in the Trees: Woody Vines of the Temperate Zone","article_sequence":1,"start_page":2,"end_page":12,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25574","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25e8128.jpg","volume":72,"issue_number":1,"year":2014,"series":null,"season":null,"authors":"Leicht-Young, Stacey A.","article_content":"Seeing the Lianas in the Trees: Woody Vines of the Temperate Zone Stacey A. Leicht-Young STACEY A. LEICHT-YOUNG Without a support structure to climb, this American wisteria (Wisteria frutescens, accession 1414-85) stretches laterally and spills over a rock wall in the Arboretum's Leventritt Shrub and Vine Garden. I n the forests and edge habitats of temperate North America, there is a group of woody plants that is well recognized but often overlooked by both the casual observer and scientific researcher alike. These woody plants are generally described as \"vines,\" but are more accurately called lianas. The ability of lianas to grow and climb in all directions, not just taller and wider like the better-known trees and shrubs, makes them a unique group of plants worthy of further study and appreciation. What is a Liana? In the simplest sense, lianas are woody vines. The term liana is better known from tropical climates where they are more abundant. By def- Lianas 3 STACEY A. LEICHT-YOUNG Lianas of the North Temperate Zone The highest diversity of liana species is in tropical areas where they can make up 25% or more of the total plant species in some forests. Lianas are much less prevalent in temperate North America, though; one estimate from North and South Carolina indicated that lianas constituted just 1.3% of the native plant species (Gentry 1991). Europe has even fewer native lianas than North America. However, southern temperate areas, such as southern South America and Eastern Asia have a higher diversity of species because of differing climate and different evolutionary history. For example, the genus Celastrus has only one native representative from North America (American bittersweet, Celastrus scandens) while China has at least 25 species NANCY ROSE Virginia creeper (Parthenocissus quinquefolia), a common North American liana, climbing up a tree trunk. inition lianas (and herbaceous vines) are plants unable to support themselves; to grow upwards, they require other plants or structures to support them. The advantage to using other plants for support is that lianas can invest resources into growing a large leaf area for photosynthesis without investing much into stem materials. A disadvantage is that when the support a liana is growing on falls down, it will also fall. However, because of their unique stem anatomy and elastic growth, they can most often resprout from their stems or roots, or simply grow along the ground until they encounter a new support. This flexibile growth habit is perhaps the defining element of lianas. The liana growth form is found in many different plant families, indicating that the climbing habit has evolved several different times. The result is a great diversity of liana species that grow worldwide in varied habitats. Celastrus angulatus is a bittersweet species from China with large leaves. MANY WAYS TO CLIMB A TREE One of the most fascinating aspects of lianas (and herbaceous vines) is the many different methods by which they can climb trees, trellises, and even walls or rock faces. In fact, Charles Darwin was one of the first to publish on the many different mechanisms that vines use to climb objects (Darwin 1867). Although there is some variation in how these groupings are made, the general categories are root climbers, adhesive tendrils, tendrils, stem twiners, and petiole climbers. Root Climbers Root climbing lianas use adhesive adventitious roots to climb trees or rock faces. These roots can often look like bunches of hairs along the liana stems. These species grow close to the substrate they are attached to and sometimes form lateral branches that grow out and away from the main stem of the liana. Familiar temperate root climbing species include poison ivy (Toxicodendron radicans), trumpet creeper (Campsis radicans), climbing hydrangea (Hydrangea anomala ssp. petiolaris), woodvamp (Decumaria barbara), and the evergreens English ivy (Hedera helix) and wintercreeper (Euonymus fortunei). STACEY A. LEICHT-YOUNG JOSEPH LAFOREST, UNIVERSITY OF GEORGIA, BUGWOOD.ORG (Far left) Hairlike aerial roots of poison ivy attach the vine to the tree. (Left) The shiny, light green foliage of woodvamp (Decumaria barbara), a rootclimbing species native to the southeastern United States. Adhesive Tendrils Like root climbers, lianas that have adhesive tendrils adhere to the tree or surface that they are climbing. However, it is not the roots that are doing the climbing in this case, but modified tendrils that have small adhesive pads at the tips. Adhesive tendril climbing lianas include Virginia creeper (Parthenocissus quinquefolia), which is one of the most common lianas in the forests of the Eastern United States; its relative, Boston or Japanese ivy (P. tricuspidata); and the showy-flowered crossvine (Bignonia capreolata), a species native to the southeastern and south central United States. STACEY A. LEICHT-YOUNG STACEY A. LEICHT-YOUNG STACEY A. LEICHT-YOUNG (Left to right) Tendrils tipped with adhesive discs cling directly to supports; flowers of a crossvine cultivar (Bignonia capreolata `Tangerine Beauty'); the unique leaves and adhesive tendrils of a wild crossvine climbing a white pine (Pinus strobus). STACEY A. LEICHT-YOUNG Tendrils are structures that are formed through modifications of the stem, leaves, leaf tips, or stipules (outgrowths at the base of a leaf). Tendrils coil around small objects such as twigs, allowing the liana to climb. The most familiar temperate lianas that use tendrils are grapes (Vitis spp.) and porcelainberry (Ampelopsis brevipedunculata), another member of the grape family (Vitaceae). Greenbrier (Smilax rotundifolia) and other Smilax species use tendrils that are actually modified thorns to climb. Although members of the genus Smilax do not technically form woody stems (they are monocots, like lilies), they are often considered to be lianas because their stems persist overwinter and form leaves in the spring. NANCY ROSE (Left) Grape tendril. (Above) Crimson gloryvine (Vitis coignetiae) is grown as an ornamental for its red to purple fall foliage. Stem Twiners Stem twining lianas, as the name describes, use their stems to climb up objects by twining around them. They can also form somewhat selfsupporting columns when many stems entwine. Stem twiners include bittersweets (Celastrus spp.) , vine honeysuckles (Lonicera spp.), wisterias (Wisteria spp.), chocolate vine (Akebia quinata), and supple-jack (Berchemia scandens), a lesser known native liana from the southeastern United States. Another species, the aromatic Chinese magnolia vine (Schisandra chinensis), is a stem twiner from one of the more ancient groups of flowering plants. Twining vines wrap around supports or even their own stems to climb. At left, entwined Oriental bittersweet (Celastrus orbiculatus) and Dutchman's pipe (Aristolochia macrophylla). Twining climbers include vining honeysuckles such as Lonicera 6 Arnoldia 72\/1 Lianas 7 STACEY A. LEICHT-YOUNG STACEY A. LEICHT-YOUNG Grape (Vitis sp., stone walls in gardens. Tendril climbers, stem far left) climbtwiners, and petiole climbers all need smaller ing on American supports to climb on since the stems or tenbeech (Fagus grandrils can only wrap around smaller diameter difolia) in mature objects such as twigs. These species are most forest. Oriental commonly observed in open forested habitats bittersweet (light or along forest edges where there are small bark) using grape (dark bark) as a supports (e.g., shrubs and small trees) and ladder to reach higher light availability. the canopy. However, some of these species--most notably grapes and Oriental bittersweet--can lianas are more abundant in high light, disemploy other methods to reach the canopy in turbed habitats because of the higher availabilolder forests with larger trees. Grapes often ity of small supports to climb on, they can be attach to trees when they are younger and conpresent in old-growth forests as well (Leichttinue to grow with them as the trees get taller, Young et al. 2010). spreading across the canopy by means of their tendrils. This is why on a walk in the woods North American Lianas and one can see very large grape stems scaling a Their Asian Relatives tree straight from the forest floor to the canopy. The liana floras of North America and East Oriental bittersweet, on the other hand, can Asia have many genera in common. For examclimb other lianas such as grapes to reach the ple, Wisteria, Clematis, Celastrus, Vitis, and canopy (this is called \"laddering\"), or it can Lonicera all have Asian and North American \"sit and wait\" in the forest understory, growing along the ground until a gap forms from a species, but Asia has greater species diversity. tree fall, resulting in higher light and smaller Since North America and East Asia share simidiameter trees growing in the gap that it can lar latitudes, many liana species (and tree, shrub, climb (Leicht and Silander 2006). So, although and herbaceous species as well) were brought 8 Arnoldia 72\/1 ROBIN BARANOWSKI STACEY A. LEICHT-YOUNG Damage to tree trunk from Oriental bittersweet. Porcelainberry (Ampelopsis brevipedunculata), originally cultivated for its attractive multi-hued fruit, has escaped cultivation through bird dispersal of seeds and is now highly invasive in edge habitats throughout much of the Northeast and Mid-Atlantic regions. STACEY A. LEICHT-YOUNG Japanese hydrangea vine cultivars (Schizophragma hydrangeoides `Roseum [left] and `Moonlight' [right]) cling to rock walls in the Leventritt Shrub and Vine Garden. ROBIN BARANOWSKI 10 Arnoldia 72\/1 Lianas 11 lenburg et al. 1993). In addition, in seasonally dry tropical forests liana roots are able to tap deep water sources over a wide area, which allows them to continue to grow during drought while trees and shrubs often go dormant (Schnitzer 2005). From what we know about species like Oriental bittersweet, they can form extensive root networks that can compete with neighboring species and contribute to vegetative spread. Thus, roots likely contribute an important part in how lianas are able to successfully colonize and persist in competition with other plant species. Intense competition from lianas above and below ground in high light situations, such as gaps in forests, may result in \"liana tangles.\" These liana tangles can suppress the ability of trees to regrow into a forest gap or slow the succession of old fields to forests for many years. In temperate areas where the growing season is restricted to the warmer months, regrowth of trees and other species may be slowed for even longer. Additionally, as lianas grow up trees they put additional stress on them, resulting in a higher chance of tree fall. This cycle of lianas increasing the chance of tree fall and resprouting in newly formed gaps may have an important influence on the regrowth A tangle of wild grape (Vitis riparia) and Oriental bittersweet climbs trees in the of subsequent secondary forests, Arboretum's Bussey Brook Meadow. especially after high-wind events ate forests. We know from tropical studies and or ice storms. These concepts have been studied a handful of temperate studies that lianas comto some extent in the tropics but need further pete with trees, not just in the obvious comobservation and research in temperate habipetition for light above ground, but also in the tats to increase understanding of how lianas commonly overlooked root zone. In temperate contribute to the composition, structure, and species, researchers have found trees competecosystem dynamics of temperate forests and ing with liana roots show slower growth rates what their future contribution may be in light than those just competing above ground (Dilof global climate change. NANCY ROSE 12 Arnoldia 72\/1 "},{"has_event_date":0,"type":"arnoldia","title":"The Pawpaw, a Forgotten North American Fruit Tree","article_sequence":2,"start_page":13,"end_page":23,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25575","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25e816d.jpg","volume":72,"issue_number":1,"year":2014,"series":null,"season":null,"authors":"Hormaza, Jose I.","article_content":"The Pawpaw, a Forgotten North American Fruit Tree Jos Pawpaw 15 savages call assemina. The French have given it an impertinent name. There are people who would not like it, but I find it very good. They have five or six nuclei [seeds] inside which are as big as marsh beans, and of about the same shape. I ate, one day, sixty of them, big and little. This fruit does not ripen till October, like the medlars.\" In 1709, John Lawson, a British explorer, reported in his book A New Voyage to Carolina--probably the first report of pawpaw in English--that \"The Papau is not a large tree. I think I never saw one a foot through; but has the broadest leaf of any tree in the Woods, and bears an apple about the bigness of a hen's egg, yellow, soft, and as sweet as anything can well be. They [the Indians] make rare puddings of this fruit\" (Lawson 1709). English naturalist Mark Catesby described and illustrated the pawpaw in his classic 1754 edition of The Natural History of Carolina, Florida, and the Bahama Islands: \"The trunks of these trees are seldom bigger than the small of a man's Leg, and are about ten or twelve feet high, having a smooth greenish brown Bark. In March when the leaves begin to sprout, its blossoms appear, consisting each of six greenish white petals, the fruit grows in clusters of three, and sometimes four together; they are at first green, and when ripe yellow, covered with a thin smooth skin, which contains a yellow pulp, of a sweet luscious taste; in the middle of which lye in two rows, twelve seeds divided by so many thin membranes. All parts of the tree have a rank, if not a foetid smell\" (Catesby 1754). In 1749, the Jesuit priest Joseph de Bonnecamps described the pawpaw: \"Now that I am on the subject of trees, I will tell you something of the assiminetree, and of that which is called the lentil-tree. The 1st is a shrub, the fruit of which is oval in shape, and a little larger than a bustard's egg; its substance is white and spongy, and becomes yellow when the fruit is ripe. It contains two or three kernels, large and flat like the garden bean. They have each their special cell. The fruits grow ordinarily in pairs, and are suspended on the same stalk. The French have given it a name which is not very refined, Testiculi asini. This is a delicate morsel for the savages and the Canadians; as for me, I have found it of an unendurable insipidity\" (Thwaites 1899). Besides these early reports, it is known that George Washington planted pawpaws at this home, Mount Vernon, in Virginia (Washington 1785). Pawpaws were also among the many plants that Thomas Jefferson cultivated at Monticello, his home in Virginia (Betts et al. 1986); during his time as Minister to France he had pawpaw seeds (Jefferson 1786) and plants (Jefferson 1787) shipped to his friends in Europe. In September 1806, the members of the Lewis and Clark expedition subsisted almost entirely on wild pawpaws for several days. William Clark wrote in his journal : \"Our party entirely out of provisions. Subsisting on poppaws. We divide the buiskit which amount to nearly one buisket per man, this in addition to the poppaws is to last us down to the Settlement's which is 150 miles. The party appear perfectly contented and tell us that they can live very well on the pappaws\" (Lewis and Clark 1806). Daniel Boone and Mark Twain were also reported to have been pawpaw fans (Pomper and Layne 2005), and early settlers also depended partially on pawpaw fruits to sustain them in times of crop failure (Peterson 1991). Pawpaws are well established in American folklore and history (the traditional American children's song, \"Way down yonder in the pawpaw patch,\" is still popular) and several towns, creeks, and rivers have been named after this fruit tree. Taxonomy, Origin, and Dissemination The first fossils of Asimina have been dated to the Eocene (about 56 to 34 million years ago) and the first clearly resembling A. triloba to the Miocene (about 23 to 5.3 million years ago) (Berry 1916). Janzen and Martin (1982) hypothesized that large fruits produced by some Central American plant species were dispersed by large mammals that were extinct by the end of the Pleistocene; they extrapolated this observation to North American plants that produce large fruits, such as the pawpaw. With the extinction of the fruit-eating megafauna, the range Naturalist Mark Catesby used flower and fruit specimens preserved in alcohol to create his illustration of pawpaw, which may explain the lack of maroon flower coloration in this depiction. Image courtesy Missouri Botanical Garden, www.botanicus.org 16 Arnoldia 72\/1 Pawpaw 17 18 Arnoldia 72\/1 Pawpaw 19 NANCY ROSE ranging from New York, and southern Michigan on the north, south to northern Florida, and west to eastern Texas, Nebraska, and Kansas (Callaway 1990). It is also present in Ontario, Canada (Fox 2012). A Description of Pawpaw The pawpaw is the only species in the Asimina genus that produces fruits of significant interest as a food source. It is, in fact, the largest edible fruit native to North America. It grows wild as a deciduous understory tree in hardwood forests with moist but well-drained and fertile soils in the eastern United States, often in large patches of the same genotype due to extensive root suckering (Kral 1960; Pomper and Layne 2005), although sometimes different genotypes can be found in the same patch (Pomper et al. 2009). Pawpaw trees can reach up to 10 meters (32.8 feet) tall and typically have a pyramidal habit in sunny locations. Pawpaw can be grown successfully in USDA plant hardiness zones 5 through 8 (average annual minimum temperatures -20 to 20 20 Arnoldia 72\/1 Pawpaw 21 MEGAN MCCARTY The Current State of the Pawpaw There was an increased interest in growing pawpaw as a crop at the beginning of the twentieth century; for example, in 1916 the American Genetic Association offered a $100 prize--$50 for the largest individual pawpaw tree and $50 for the tree--regardless of size--with the best fruit (American Genetic Association 1916). Yet in spite of its high potential through the years as a new high-value niche fruit crop, pawpaw is still only in the early stages of commercial production. The greatest current market potential for pawpaw is probably in local markets and direct sales to restaurants and other gourmet niche customers. Most pawpaw fruits are A female zebra swallowtail oviposits on an emerging pawpaw leaf. Butterflies, Pawpaw, and Coevolution The zebra swallowtail (Protographium marcellus, formerly Eurytides marcellus) is a beautiful black and white striped butterfly whose caterpillars feed exclusively on Asimina leaves. (The damage made to the leaves is reported to be negligible in pawpaw orchards [Pomper and Layne 2008]). Some compounds present in the pawpaw leaves (acetogenins, specific substances only found in species of the Annonaceae) are repellent to most insects and birds so the caterpillar accumulates them to avoid predation. These natural bioactive compounds present in the leaves, bark, and twigs of pawpaw and other species of the Annonaceae have shown some insecticidal and anti-tumoral properties (McLaughlin 2008). Of the seven swallowtail tribes, the Graphinii (to which the zebra swallowtail belongs) is one of the largest with about 150 species restricted to the tropics and subtropics except for two, Iphiclides podalirius and Protographium marcellus (the zebra swallowtail), that live in Palearctic and Nearctic regions, respectively (Haribal and Feeny 1998). The fact that both the zebra swallowtail and the pawpaw are the only members of their respective groups to live in temperate North America indicates that both species have coevolved and provides a neat system to study coevolution and adaptation to cooler climates. Illustration of the exterior and interior of a pawpaw fruit painted by Royal Charles Steadman in 1924. From the USDA Pomological Watercolor Collection in the Rare and Special Collections of the National Agricultural Library in Beltsville, Maryland. 22 Arnoldia 72\/1 Pawpaw 23 Jefferson, T. 1786. From Thomas Jefferson to John Bartram, with Enclosure, 27 January 1786, Founders Online, National Archives (http:\/\/founders.archives.gov\/documents\/ Jefferson\/01-09-02-0201, ver. 2013-08-02). Source: The Papers of Thomas Jefferson, vol. 9, 1 November 1785 "},{"has_event_date":0,"type":"arnoldia","title":"Biodiversity Hotspot: China's Hengduan Mountains","article_sequence":3,"start_page":24,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25573","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25ebb6f.jpg","volume":72,"issue_number":1,"year":2014,"series":null,"season":null,"authors":"Boufford, David E.","article_content":"Biodiversity Hotspot: China's Hengduan Mountains David E. Boufford I n southwestern China, in the southeastern corner of the Qinghai-Tibet Plateau, lies one of the world's 35 biodiversity hotspots: the Hengduan Mountains. This hotspot occurs at the juncture of mountain systems where precipitation can vary tremendously due to a combination of topography, climate, and hydrology. The terrain forms topographic channels that funnel seasonal monsoon rains up through the river valleys from the lowland tropics of south- ern China, India, and Myanmar (Burma) to the southeastern edge of the 5,000-plus-meter-high (16,400-plus-feet) Qinghai-Tibet Plateau. The region also receives vast amounts of water from the five major rivers that drain the plateau: the Yarlung Zangbo Jiang (which becomes the Brahmaputra in India and the Jamuna in Bangladesh); the Ayayerwaddy (Irrawaddy); Nu Jiang (Salween); Lancang Jiang (Mekong); and Jinsha Jiang (known in the West as the Yangtze, and Qinghai-Tibetan Plateau Nepal Bhutan Myanmar Hengduan Hotspot India Hengduan Hotspot 25 ALL IMAGES BY THE AUTHOR Sichuan, Batang Xian. A glacial lake lies at about 4,500 meters (14,764 feet) on the south side of the pass at Haizi Shan, surrounded by a Kobresia (bog sedge) meadow and with scattered dwarf Salix and Rhododendron on nearby slopes. What's a Biodiversity Hotspot? As defined by Conservation International, to qualify as a biodiversity hotspot a region must meet two strict criteria: 26 Arnoldia 72\/1 Hengduan Hotspot 27 Sichuan, Litang Xian. North of Litang between Litang and Xinlong. A broad ravine with numerous side seepages and both moist and dry upland meadows, featuring the tall, yellow-bracted floral spikes of Rheum alexandrae (a rhubarb relative) and yellow-flowered Pedicularis longiflora var. tubiformis in the foreground along the stream. centron, Acer (maple--45 species!), Aesculus (buckeye), Tilia (linden), several genera within Lauraceae (the laurel family), Meliosma, Phellodendron (corktree), Evodia, Cornus (dogwood), Ostryopsis, Carpinus (hornbeam), Ostrya (hophornbeam), Betula (birch), Quercus (oak), Lithocarpus, Fagus (beech), Elaeocarpus, and Ailanthus (Boufford and Ohba 1998). In formerly glaciated valleys and on higher slopes, Abies (fir), Picea (spruce), Betula and other boreal plants intermix with vegetation generally considered to be warm-temperate. Full grown, well-formed oak and fir trees reach an elevation of around 4,600 meters (15,092 feet) in some places and intermix with alpine meadows, scree slopes, and Rhododendron thickets. Herbaceous vegetation reaches to 6,000 meters (19,685 feet), although few plant specimens have been collected above 5,500 meters (18,045 feet). The east and southeast portions of the Hengduan region are the best known, since they were easily reached by explorers and researchers coming up the Chang Jiang (Yangtze) or entering from Chengdu, about 100 kilometers (62 miles) away, or from Kunming. The forests on the eastern slopes also harbor some of the last surviving populations of giant panda, and China's best known and perhaps largest panda research station at Wolong Shan. About half 28 Arnoldia 72\/1 Hengduan Hotspot 29 Wang Qia photographing plants at Haizi Shan, an extensive cold, glaciated plateau with numerous lakes, ponds, and streams that often flow out of sight under the glacial debris. up through the middle of shrubs, which offer some protection from the animals. The reason that so many herbarium specimens of herbaceous plants from these areas lack underground organs is because of the difficulty in extracting them from the middle of the coarse, frequently spiny shrubs in which they grow. dendron (226 species), Pedicularis (217), Saussurea (100+), Ligularia (70), Cremanthodium (38), Anaphalis (33), Leontopodium (25), Artemisia (55), Gentiana (117), Primula (113) Saxifraga (136), Salix (103), and Corydalis (89). EARLY EXPLORATION The first western explorers (and essentially the first naturalists) in the area were French missionaries who traveled to the remotest regions of China to convert the locals to Christianity (Kilpatrick 2014). Most were also trained in the natural sciences and were encouraged to collect and send specimens back to Paris. Among the most notable of these missionaries were P 30 Arnoldia 72\/1 Sichuan, Baiyu Xian, Zhandu Xiang. Mixed conifer-mixed broadleaved deciduous forests, with Salix (willow) growing near the stream, along a branch of the Ou Qu in Ase Gou (Ase Gorge). References Cited Boufford, D. E. and P. P. van Dijk. 2000. South-Central China. pp. 338 32 Arnoldia 72\/1 Hengduan Hotspot 33 GENTIANACEAE Gentians (Gentiana) and their relatives, notable for blue flower color, are characteristic of the Hengduan flora. Of the 248 species of Gentiana in China, 117 occur in the Hengduan region. Clockwise from upper left: 34 Arnoldia 72\/1 Hengduan Hotspot 35 PEDICULARIS Pedicularis has its main center of distribution in the Hengduan Mountain region, where 217 of China's 352 species occur. The plants are reported to be hemiparasitic, getting at least part of their nutritional needs from host plants. Despite many attempts, we have been unsuccessful in excavating the underground parts to find the connections with a host plant. While easy to recognize vegetatively as being a species of Pedicularis, the flowers are needed for identification to the species level. The beak (galea) and orientation of the corolla appear to be correlated with pollination by bumblebees (Eaton et al. 2012; Wang and Li 2005). The Hengduan region, with 65 species, is also the world's center of diversity for bumblebees (Williams et al. 2009). Clockwise from upper left: "},{"has_event_date":0,"type":"arnoldia","title":"A Shady Character: Platanus x acerifolia","article_sequence":4,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25572","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25eb76b.jpg","volume":72,"issue_number":1,"year":2014,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"A Shady Character: Platanus "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23440","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170b328.jpg","title":"2014-72-1","volume":72,"issue_number":1,"year":2014,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"John George Jack: Dendrologist, Educator, Plant Explorer","article_sequence":1,"start_page":2,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25570","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25eb36d.jpg","volume":71,"issue_number":4,"year":2014,"series":null,"season":null,"authors":"Pearson, Lisa","article_content":"Lisa Pearson J ohn George Jack was a notable figure in the early history of the Arnold Arboretum. His story is perhaps less well known than those of his colleagues, but his fifty-year dedication to the study of trees, plant exploration, formal and informal education, and especially the instruction of a generation of Chinese botanists is unmatched. In 2012, the Arboretum was fortunate to acquire a trove of John Jack archival materials from his granddaughter, Constance W. Cross. Included were three manuscripts written by Jack in the early 1940s, in which he gives a lively account of his early life in Canada as well as a detailed look at the beginnings of the Arnold Arboretum. These historical sketches provide new insight on Jack and served as a primary resource for this article. Roots John George Jack (1861 John George Jack 3 Troy Female Seminary in Troy, New York, and taught in the city for several years before moving to Quebec. She had been the teacher at the Protestant school in Ch 4 Arnoldia 71\/4 John George Jack 5 continued in the community and were taught by many of the same people who had originally taken classes from Jack years before. In the first twenty years of his career in Boston, John Jack was a bachelor of presumably thrifty habits, boarding with Mrs. Cheney in Jamaica Plain. As such, he was able to accumulate enough savings to periodically travel in order to botanize and visit botanic gardens and arboreta. Travel in that period was truly an expedition; it took about a week to reach Europe by sea, and once there, ground transportation was by rail or horse-drawn conveyance. Jack's trips were lengthy, lasting six months in the case of his visit to Asia (see textbox on page 6). He made his first trip overseas in 1891, visiting Paris, Berlin, Geneva, northern Italy, Copenhagen, Hamburg, Brussels, and Britain. He spent several weeks at Kew alone and at An announcement for a spring session of John Jack's popular tree classes from around 1900. every stop was able to meet in person the botanists and horticulturists with whom he had corresponded. He took another leave of absence in 1898 to explore and report on the forests of the Pikes Peak region, his first exposure to the Rocky Mountain flora. Jack went west again in 1900 to This image of Abies lasiocarpa at Lake Louise, Alberta, Canada, was made by Alfred Rehder in August 1904 while on a plant collecting trip with John Jack. A JOURNEY TO THE FAR EAST In 1905, John Jack decided to visit Japan, Korea, and China. He hoped that the things he would invariably learn while abroad and the plants he might find would enrich his teaching and the collections of the Arboretum. For some unknown reason, Charles Sargent was opposed to his trip. He refused to pay for any of Jack's expenses and he docked Jack's pay of fifty dollars a month for the duration of his sixmonth leave of absence. Undeterred, Jack left Boston at the beginning of July and arrived in Yokohama at the end of the month. He spent the next month and a half visiting gardens, parks, and forests in the area and made an expedition further afield to Nikko and Lake Chuzenji. He decided to alter his itinerary and pay a visit to Sapporo where he was hosted by Professor Kingo Miyabe, whom he had known many years earlier when Miyabe was a doctoral candidate at Harvard University. From Japan, Jack sailed to Korea where he spent several weeks exploring the region around Seoul. Unfortunately the Japanese government, which had ruled the country since the end of the recently concluded Russo-Japanese War, would not allow travel out of the area, thus precluding any chance of botanical collections outside of the capitol. Jack then traveled to Shandong, China, and then on to Beijing. There he spent time botanizing with his old friend Frank N. Meyer, who was collecting economic plants for the U.S. Department of Agriculture. He returned to Japan in October to spend time with his brother, Milton, and to revisit Lake Chuzenji where he had noted numerous rhododendron and azalea species from which he collected seeds. He finally sailed for home by way of Naples, Italy, in November, arriving in New York on December 20. Jack considered this trip a success, notwithstanding the recently concluded war between Russia and Japan that hampered his movements somewhat. It cost him some $2,000, so it came as a pleasant surprise when Sargent, in an uncharacteristically apologetic manner, admitted the great value of Jack's collections and allowed him the $300 in back pay that had been withheld during the trip. The library has digitized a collection of John Jack's photographs from Japan, which are available at http:\/\/via.harvard.edu. A short introduction is available on the Library website: http:\/\/arboretum. harvard.edu\/library\/image-collection\/botanical-and-cultural-images-of-eastern-asia\/john-george-jack\/ Japanese black pine (Pinus thunbergii) grows above the wall and moat surrounding the Imperial Palace in Tokyo, Japan, in this John Jack photo from August 19, 1905. John Jack photographed this large specimen of Japanese chestnut (Castanea crenata, known then as C. japonica) along a road between Narai and Fukusawa, Japan, on September 2, 1905. In addition to making his own photographs in Asia, Jack also purchased colored lantern slides to use in his lectures. Seen here are two lantern slides from Japan showing people under a wisteria-covered arbor (left) and women digging shellfish on a beach (right). 8 Arnoldia 71\/4 John George Jack 9 could matriculate at the Bussey Institution and then study with John Jack, one-on-one or in small groups. An early student was WoonYoung Chun (Chen Huanyong) who had previously studied forestry at the Massachusetts Agricultural College and the forestry school at Syracuse University and came to study dendrology with Jack in 1915. Students like Chun came halfway around the world to study the tree flora of their native country because of the convenience of having an extensive living collection and a complete herbarium all in one place. In a 1917 interview Chun remarked, \"It would take me a lifetime of travel to study what I can find out here about Chinese trees in a few years.\" One of Jack's most notable Chinese students was H. H. Hu (Hu Xainsu), the botanist who, along with colleague W. C. Cheng (Zheng Wanjun), first identified and named living examples of dawn redwood (Metasequoia glyptostroboides), a tree previously thought to be extinct but found growing in Hubei in the late 1940s. Hu greatly respected and admired Jack and corresponded with him for the remainder of his life. In a letter dated June 17, 1931, Hu asks Jack for a portrait that they might hang in their herbarium, \"Since most of Chinese systematists studied under you and you have exerted such an important influence toward Chinese botany, your photograph is specially needed.\" In addition to their education at the Arboretum, Jack also brought his students to his property, \"Folly Farm,\" in Walpole, Massachusetts, for practical horticultural training in the garden and orchard. John Jack (far left) with some of his Chinese students, including Woon-Young Chun to the far right in the boater hat, in a 1917 photograph by A. A. Greenlaw. ARNOLD ARBORETUM JACK'S ACCESSIONS John George Jack left an indelible mark on the Arnold Arboretum, particularly through the prudent care and attention he gave to the early curation of the rapidly expanding collection. He was also quite the collector of plants himself and over 1,700 accessions originally collected by Jack have moved through the Arboretum. These represent collecting efforts in Asia as well as considerable sampling throughout North America. The majority of his collections did not survive beyond the 1930s, but some 100 accessioned plants collected by Jack do continue to grow at the Arboretum. These include three interesting hybrids, all named in honor of Jack, and some Korean accessions from Jack's 1905 trip to Asia. John George Jack 11 Legacy In 1926, Charles Sargent personally asked Jack to go to the Atkins Institution in Cuba, near Cienfuegos in the western part of the island, to collect specimens for the Arboretum herbarium, which lacked material from that part of the Caribbean. He made several trips over the next ten years, sometimes accompanied by special students from the Arboretum. The Atkins Institution was started as a private experiment station at about the turn of the twentieth century to develop better varieties of sugar cane. It was given to Harvard some years later and comprised over two hundred acres of open and forested land populated with Cuban and West Indian woody plants. In addition to his collections for the Arboretum, Jack also began a herbarium for the use of the Institution containing specimens from their collection as well as other Cuban flora. Karl Sax, Bussey Institution colleague and future director of the Arnold Arboretum, spent time with Jack at the Atkins Institution in 1936. He remembered, \"I discovered that although he was 75 years old Professor Jack was up at 6 A.M., worked all day, often travelling into the surrounding country on horseback, and continued to work until 11 or 12 o'clock at night.\" The Institution remained part of Harvard until 1961 when its director, Dr. Duncan Clement, left Cuba due to pressures associated with the Cuban Revolution and the University ended its support. Today the garden is managed by the Cuban government. John Jack continued to busily curate the Arboretum collections, to teach, and to collect plant material up until his retirement in 1935 at age 74, the mandatory retirement age imposed by Harvard University. In his later years he lived on his farm in Walpole where he maintained an extensive apple orchard that yielded large crops every year. His wife Cerise had died just after his retirement but his adopted daughter Betty, her husband, and their two daughters shared the farm with him. While pruning in his orchard in 1948, Jack fell from a ladder and broke his hip, leaving him bedridden. He died several months later in 1949, aged 88. A person like John George Jack would be a rarity today. He was a smart self-starter who made the most of opportunities when they pre- John Jack in Cuba, April 1927. sented themselves, and was fortunate to live at a time when it was not absolutely necessary for an academic to have an advanced degree. He was a teacher with an amazing gift for engaging his students, no matter what their background and education might have been. He was a methodical and diligent naturalist, in the broadest meaning of the term, whose interests ranged from entomology to forestry, horticulture, dendrology, and all points in between. The Arnold Arboretum was extremely fortunate that Jack chose to spend his long career here in Boston. Acknowledgement We would like to extend our sincerest thanks to Constance W. Cross for her generous gift of John Jack archival materials. Lisa Pearson is Head of Library and Archives at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Book Excerpt-Hemlock:A Forest Giant on the Edge","article_sequence":2,"start_page":12,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25568","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25eaf6f.jpg","volume":71,"issue_number":4,"year":2014,"series":null,"season":null,"authors":"Foster, David R.","article_content":"BOOK EXCERPT Hemlock: A Forest Giant on the Edge David R. Foster, Editor Editor's Note: Eastern hemlock (Tsuga canadensis) is an iconic tree species in northeastern forests and the Appalachian Mountains. It has faced peril in the past but is now faced with perhaps its most deadly threat--the invasive and devastating insect pest, hemlock woolly adelgid. In this new book, Harvard Forest director David Foster and several colleagues and scientific collaborators explore the history and ecology of and challenges to the majestic eastern hemlock. Presented here by permission of the publisher is an excerpt from Chapter Three: Prehistory to Present, written by Wyatt Oswald, David Foster, and Jonathan Thompson. In the previous part of the chapter the authors describe the process of extracting 3-inch-wide, 3-foot-long sediment cores from a pond for later paleoecological analyses of the material. Hemlock: A Forest Giant on the Edge David R. Foster, Editor. Written by Anthony D'Amato, Benjamin Baiser, Aaron M. Ellison, David Foster, David Orwig, Wyatt Oswald, Audrey Barker Plotkin, and Jonathan Thompson; Stephen Long, Consulting Editor. Yale University Press. 2014. 336 pages. ISBN: 978-0-300-17938-5 Book Excerpt 13 H EMLOCK has changed in abundance numerous times in the past, and it now faces an extreme threat from the hemlock woolly adelgid. As we seek to consider this new dynamic in perspective, we are fortunate that hemlock has left a remarkable array of records that shed light on its ecology under a wide range of conditions. These historical and paleoecological archives inform the field studies, experiments, and modeling activity that we undertake in the woods and back in the laboratory. A look at hemlock's fossil record helps us examine how hemlock has changed with the intense human activity in the past few centuries and allows us to assess how it might cope with the combination of insect onslaught, climate change, and ongoing human activity today and in the future. It also enables us to evaluate whether there is any hope that hemlock may stave off or recover from the population collapse associated with a new invasive organism. We use a variety of tools and techniques to reconstruct the historical dynamics of the forest environment and vegetation, as well as individual tree species. To reach back furthest, we study pollen, other microscopic fossils, and diverse signatures of past environments that are preserved for millennia in the sediments of lakes, bogs, swamps, and other wetlands. More recent centuries and decades come alive in historical land-survey documents, field studies of old-growth forests, and tree rings that yield insights into the composition and structure of forest vegetation from the time of European arrival forward. In some cases, the particular qualities of hemlock provide a record that bridges prehistory and history. For example, by carefully dissecting the deep beds of needles that accumulate on the cool, moist ground beneath hemlock, we find pollen and other plant parts that yield a chronological record connecting the postglacial period with the time since European settlement. From these distinctive soil layers comes a record of changes in the composition of individual forest stands that can be linked to the evidence from tree rings, uprooted trees, and the many other clues that are present in the hemlock forest itself. Those of us conducting retrospective studies at the Harvard Forest have employed this full array of approaches, exploiting every opportunity to reconstruct the distribution, abundance, and dynamics of hemlock across New England and going back thousands of years into the past. ONE HUNDRED and fifty years ago, Henry Thoreau mused in his journal on what stories might be gleaned from the pollen grains accumulating in small pools and ponds, but it took nearly a half century more for the Swedish naturalist and geologist Lennart von Post to first take advantage of this phenomenon in studying the history of plants over long periods. He published a report in 1917 showing that the grains of pollen identified in Scandinavian peats told an astonishing story of dynamic changes in vegetation composition. Two characteristics of pollen make it a particularly useful tool for interpreting the past. First, pollen grains are remarkably durable because they are shielded by an outer layer of complex chemical compounds that protect the sperms cells as they get transferred from the stamens to the pistils of flowering plants, or from male to female cones in conifers like pine or hemlock. Second, the pollen of different species and genera of plants is different enough to allow us to identify them. It comes in a wide range of shapes, sizes, and surface markings, all of which allow palynologists--the meticulous and patient scientists 14 Arnoldia 71\/4 Book Excerpt 15 ALL IMAGES BY DAVID FOSTER The top of an eastern hemlock (Tsuga canadensis) pokes above the canopy on the Prospect Hill tract of Harvard Forest. Researchers extract sediment cores from Harvard Forest's Hemlock Hollow. An extracted sediment core is finished and labeled by researchers. 16 Arnoldia 71\/4 Book Excerpt 17 previous generations of plants. In New England, where glaciers scoured the earth surface during the last ice age, the duration of both of these sedimentary archives is limited to the period since the ice melted, the land surface stabilized, and the climate allowed the growth of plants. Thus, the oldest lake records span about twelve to fifteen thousand years, and many wetlands only extend back five or six thousand years. MEANWHILE, back in the lab, we slice the cores into thin sections, half an inch or less in length, and carry out a series of treatments and analyses of the material. It's not just pollen grains that we seek. For instance, we want to know the age of the mud at different depths in the core, so we extract small samples of sediment or plant material and send them to a specialized (and expensive) laboratory that assesses the radiocarbon content of the material. We also measure the sediment's organic and mineral content or particle sizes to determine changes in the lake environment, including past droughts, which are often registered as layers of sandy, inorganic material. In combination with other chemical analyses, these sedimentary characteristics provide a detailed record of past variations in climate. We isolate pollen grains as well as the spores from ferns and other early plants by subjecting mud samples to intense acid baths, washings, centrifuge spins, and sieving steps. It's remarkable that these intense treatments remove most of the organic and mineral material but leave a tiny residual fraction that contains the concentrated and quite intact pollen, along with bits of insects, charcoal, and other miscellaneous detritus. The tiny pieces of charcoal and insect remains, both of which are as highly resistant to decay as pollen, are sieved, identified, and counted under a microscope to provide information about past wildfire activity and insect outbreaks. We mount the residue on microscope slides and examine them with high-powered magnification, carefully scrutinizing and identifying every pollen grain that is encountered. At any given level, a palynologist might identify 300 to 500 pollen grains through a painstaking process that can take anywhere from two to eight hours or more. Pollen data tell us the relative abundance of different species. If 50 out of 500 pollen grains at a given level are identified as hemlock, this would yield a value of 10 percent. Knowing whether or not a species is a prolific pollen producer helps us to assess how well the relative abundance of its pollen corresponds to its actual abundance on the landscape. The pollen of insect-pollinated trees such as maple and chestnut rarely exceeds 5 percent of the total, whereas pine, birch, and oak can easily reach 10 to 20 percent or more. Considering these factors, we would assume that 5 percent chestnut means a significant presence. At its very crudest, a pollen diagram will show at what point in the past hemlock or any other plant was absent, rare, or abundant. In most cases, it will also reveal fascinating curves depicting the long-term variation in these species in relationship to other species and many environmental factors. In well-studied regions such as eastern North America, many dozens of pollen records have been analyzed over the last few decades. In southern and central New England, the Harvard Forest group has analyzed cores from more than three dozen sites. We make the data available to everyone electronically on our website and collaborate with many people who use them. We also keep the cores from which samples have been taken in cold storage for our future needs and those of other scientists who may be interested in examining our records in more detail or for searching for other materials and clues in the mud. Our 18 Arnoldia 71\/4 Book Excerpt 19 one of great importance today because of the looming likelihood of rapid climate change and the question of how plants will respond and cope with new conditions. We are employing all sorts of approaches--genetics, simulation modeling, field and laboratory studies of dispersal, and pollen analysis--as we continue to grapple with the question. Have we overestimated the rates at which trees moved in the past, or are we underestimating their anticipated and potential future dispersal rates? One possible way to account for a more rapid past dispersal is to invoke a history of rare long-distance dispersal events, such as abrupt gusts and updrafts in wind that may loft a seed into the jet stream, or the rare flight of a bird in which it carries a seed for dozens of miles. In this way, a chance event can disperse seeds great distances. If such an event happened even once a decade, it may have been extremely important in shaping patterns of movement over centuries. We cite uncommon processes such as these in our modeling discussions when talking about the dispersal of insects like the hemlock woolly adelgid or the adaptations of plant species under future climates. As research on this dilemma progresses, the answers to these questions will have important implications for predicting the future shape of our forest ecosystems and for gauging the ability of many species to survive the expected changes in climate in coming decades. The long-term history of hemlock also reveals the extreme malleability of forest types and assemblages, including those that are familiar to us today. Hemlock arrived in the northeastern United States about 2,000 years after white pine and 2,000 years before American beech, even though today it frequently grows alongside both these species, and we often think of them as members of the same plant communities. Given beech's similarity to hemlock in shade tolerance and suitability for forest canopies, and the manner with which they coexist in many places today, it is hard to imagine that hemlock grew in New England for 2,000 years without beech. Similarly, it was only with the arrival of hemlock that the New England landscape developed forests akin to the old-growth stands of white pine and hemlock studied by early ecologists and described in many Harvard Forest studies, including those by Richard Fisher, Bob Marshall, Tony D'Amato, and Dave Orwig. The contrasting histories of these various trees illustrate that species respond in highly individualistic ways to environmental change. Because conditions in the past were distinctly different from the present, we witness the species behaving in significantly different ways over time. The assemblages of plants and animals that are familiar to us today are actually quite ephemeral in deep time and space. It is through such understandings that we've developed an ecological theory that accepts and explains the separate though interactive behavior of species. One of the earliest and best articulations of this theory came from a noted northeastern botanist--Henry Gleason of the New York Botanical Garden--who developed the \"individualistic concept of ecology\" in the early 1900s. This simple but revolutionary theory posited that the makeup of vegetation on a site was determined by the actions of the many individual species, each of which operated quite separately from others and according to its unique ecological qualities. Although this concept was debated for decades, some of the strongest evidence that led to its conclusive support came from paleoecological studies that showed the highly disparate behaviors of different tree species in migration and in response to climate change and to natural and human disturbances. While this understanding of plant behavior and ecology emerged from the past and helps us explain our current landscapes, it should also prepare us for unanticipated combinations of species to appear under the anomalous conditions expected for the future. 20 Arnoldia 71\/4 Book Excerpt 21 Coring dozens of ponds and bogs and examining tens of thousands of pollen grains preserved in their sediments has helped us outline the following picture of New England's prehistory. After a lengthy dry period, from around 11,500 to 10,000 years ago, during which white pine dominated the landscapes of the northeastern United States, hemlock increased in abundance across much of New England, then reached its peak population levels during a relatively warm and moist interval from 8,000 to 5,500 years ago. Beech had arrived to join hemlock in the region at that point, and with oaks, birches, and maples also present, and white pine and pitch pine already well established, the overall composition of New England forests was quite similar to what we find in our landscape today. Although the environmental conditions of that earlier time appear to have been well suited for hemlock, some of our recent research suggests that brief periods of cold climate occurred every few centuries, with deleterious impacts on hemlock in some parts of New England. Various lines of evidence, including chemical analyses of lake sediment records, show that the generally warm, moist conditions were interrupted occasionally by a century or so of cold, dry climate. And while hemlock and other species did not always respond uniformly to these events across the region, some of our relatively detailed pollen records feature abrupt, short-lived declines of hemlock, including significant population reductions at around 8,000 and 6,000 years ago. Hemlock certainly didn't disappear from the landscape during these events, but the pollen data do suggest that it became much less abundant during times of cold, dry conditions. Then, around 5,500 years ago, hemlock experienced an abrupt, range-wide collapse. For about two millennia it nearly disappeared throughout its entire range in the Northeast before it rebounded about 3,500 years ago. Although it recovered greatly across the region, at most sites hemlock never returned to its predecline levels. This hemlock decline is one of the most thoroughly studied aspects of the postglacial vegetation history of North America, yet we still don't completely understand what caused it or sustained it. Conclusions drawn over the past three decades variously attribute hemlock's decline to a species-specific disease, a massive insect outbreak, a sustained shift to drier climate, a series of drought events, and a combination of these factors. It is now quite clear that climate was strongly involved and that in some ways the big decline was a larger version of the earlier declines witnessed during cold spells. If the trees weren't killed directly by drought, then the associated environmental conditions either stressed hemlock in ways that made it more susceptible to insects or disease or facilitated an unusual outbreak of a pest or pathogen. (It was this record of minor events leading to the major drought and decline in hemlock that our colleague correctly surmised he was seeing in the various layers of sand we observed that day on the raft in the middle of the lake.) Hemlock eventually recovered, and pollen records reveal that it was again abundant in New England forests from around 3,500 years ago to the time of European settlement. Our studies of the sediments of Hemlock Hollow, a vernal pool hidden in the large hemlock forest on the Prospect Hill tract of the Harvard Forest, have yielded a detailed stand-scale record of forest changes over the last 10,000 years. The local nature of this record enables us to examine the fine-scaled ecological response of an individual forest to various changes in its environment. Here we can see that when disturbances occurred, including fires every 1,000 to 3,000 years, hemlock abundance dropped abruptly and then rebounded slowly, taking 500 years or more to recover to original levels. In the recovery from these major disturbances--intense events that we interpret to have killed most of the larger trees--the 22 Arnoldia 71\/4 Book Excerpt 23 Eastern hemlocks and eastern white pines along the Swift River in Petersham, Massachusetts. 24 Arnoldia 71\/4 Book Excerpt 25 regions of the world, including glacial advances farther north. It may seem counterintuitive that two species common in northern New England would be bothered by a shift to colder climate. It is quite possible, however, that conditions became both colder and drier, with both hemlock and beech suffering due to their relatively high moisture requirements. The latter part of the Little Ice Age coincided with the expansion of European colonists across New England, transforming the land. Region-wide, up to 60 percent of the land was cleared for agriculture and the rest was cut--repeatedly in some places--with a peak in harvesting occurring in the late nineteenth and early twentieth century. Although forest once again covers more than 80 percent of New England, these second-growth stands are not the same as those of presettlement times. When we compare the witnesstree data with present-day forest composition, we find that some species are more common than they were centuries ago, such as early successional birches, red maple, and pines, including the old-field white pines that invaded abandoned agricultural lands. These light-seeded, fast-growing, and light-requiring species spread and grew rapidly across heavily disturbed areas, thriving after the intense farming and logging subsided. On the other hand, some species are less abundant than they were before European settlement. Species of mature forests, including hemlock and beech, are much less common than they were in the witness-tree surveys. Throughout the Northeast, hemlock declined as much as 10 percent over the last 400 years. When we zoom back in from the region-wide scale to that of the individual landscape, we often see considerable evidence of land use in the characteristics of hemlock forests. In some cases, seemingly ancient hemlock stands have undergone much greater changes in their recent past than we might at first assume. These are the unexpected findings of a study led by Harvard Forest researchers Jason McLachlan and David Foster. They set out to reconstruct the histories of four old hemlock forests in central Massachusetts, using both tree-ring analysis of the largest trees and centimeter-by-centimeter analyses of pollen grains preserved in the approximately six-inch-thick layer of organic matter forming the top layer of the soil. They found that the stands, dominated today by hemlocks 100 to 200 years old, had experienced a series of disturbances over the last few centuries, including logging, windstorms, fires, and pathogen outbreaks. Indeed, early and mid-successional trees such as oaks, pines, and American chestnut had occupied those same stands at different times in the past. In many of the forests, it appeared as though today's dominant hemlocks may in fact owe their current good fortune to the removal of competing species by selective logging and the chestnut blight. Like many of our other retrospective investigations of hemlock, this study of secondgrowth stands obliges us to change the way we think about the species, the forests it forms, and the way that nature operates. On one hand, forests that appear to be unchanging may be relatively recent in origin and shaped by processes that the species has never experienced before. On the other, although hemlock forests have been dynamic at times, the history of the species in New England has always been one of long-term dominance interrupted by infrequent abrupt declines. With such a decline spreading across the landscape today, we can expect another lengthy period with little hemlock followed by--we can only hope--its gradual return. Wyatt Oswald is Associate Professor at Emerson College, David Foster is Director of Harvard Forest, and Jonathan Thompson is Senior Ecologist at Harvard Forest. "},{"has_event_date":0,"type":"arnoldia","title":"Getting Buzzed at the Arnold Arboretum","article_sequence":3,"start_page":26,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25569","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25eb328.jpg","volume":71,"issue_number":4,"year":2014,"series":null,"season":null,"authors":"Switzer, Callin","article_content":"Getting Buzzed at the Arnold Arboretum Callin Switzer W hile strolling through the Arnold Arboretum during the summer, visitors may see bees flying from flower to flower. Some bees are pushing their heads deep into flowers and collecting nectar, others are more interested in collecting pollen from the flowers' anthers. Bees that collect pollen are not collecting most of it for themselves; they are taking it back to their colony to feed to the larvae. Pollen provides a protein and mineral source for the developing brood. In flowering plants, pollination is the process of moving pollen from the anthers to the stigma. As bees collect pollen and nectar, they inadvertently transfer pollen from the anthers of one flower to the stigma of another flower, either on the same plant or different plants. For plants that are self-incompatible (they cannot reproduce without cross pollination from another plant), this transfer service is essen- tial. For both self-incompatible and self-fertile species, the transfer of pollen between plants allows for genetic variation in the plants' offspring, preventing plant populations from becoming inbred. Without bees, many species would make neither fruit nor seed. Give It Up for the Bees Plants have developed a variety of ways to \"give\" their pollen to bees. Some have longitudinally dehiscent anthers; these split open down the sides when the pollen is ready, making it easily accessible. Longitudinally dehiscent anthers have benefits and drawbacks. One benefit is that plants with these anthers can be pollinated by many insects, birds, or even humans. (In Sichuan, China, the decline of pollinators has led to pollination of apple and pear orchards by humans armed with vials of pollen and small brushes.) These plants are generalists CURT RAWN Pass the Bees, Please The next time you dip into a bowl of salsa, serve up butternut squash soup, or savor a slice of blueberry pie, thank the bees. Tomatoes, squash, apples, blueberries, and lots of other delicious foods require pollination--mostly done by bees. About 30 percent of our food relies directly on pollinators, and thousands of plant species depend on bees for reproduction. Honey bees and bumblebees are major pollinators of food crops, but many other bee species also pollinate a wide range of plants. Buzz pollination helps produce plump tomatoes like these (Solanum lycopersicum `Lemon Boy'). Buzz pollination 27 The anthers of Chinese stewartia (Stewartia sinensis) (left) and Carefree Beauty rose (Rosa `Bucbi') (above) are longitudinally dehiscent, which means that each anther splits along its length to release pollen. when it comes to attracting pollinators, but this could also be seen as a drawback. What if an animal rushes by and knocks all the pollen off the flowers? What if an insect visits different species and never actually transfers pollen between conspecifics (members of the same species)? Changing the shape of the anther can help solve both of these potential problems. Multiple lineages of plants have evolved anthers that are tube-shaped. Instead of splitting down the sides, these anthers simply open tiny pores when pollen is ready to be released. These anthers are known as poricidally dehiscent, or poricidal. Poricidal anthers help keep pollen from being knocked off the flower, and they prevent many pollinators from reaching the pollen. About eight percent of flowering plants (some 20,000 species) have poricidal anthers (Buchmann 1983). Because the pollen is in a tube, animals cannot easily shake it free. Three common ways to access this pollen are biting through the outside of the anther; squeezing pollen out by treating the anther like a tube of toothpaste; or A bittersweet nightshade (Solanum dulcamara) flower, showing poricidal anthers. At the center of the flower, ten distinct pores are visible. Pollen is held in the tubes below the pores. Purple beautyberry (Callicarpa dichotoma) has poricidal anthers; a pore is visible on the lower edge of the anther on the right in this photo. ALL PHOTOS BY THE AUTHOR UNLESS OTHERWISE INDICATED 28 Arnoldia 71\/4 Buzz pollination 29 Look closely, and you can identify bumblebees and honeybees. Honeybees (left) are more slender and brown. They usually have rings of grey on their abdomen. Bumblebees (right) are round, fuzzy bees, often with black on the abdomen and yellow on the thorax. A common eastern bumblebee (Bombus impatiens) nest that was reared in a lab; the cotton covering has been pulled back to expose the cells. Nests are typically built in preexisting holes, often below ground. The structure is mostly made of wax, which is secreted from between the segments of a bumblebee's abdomen. A typical bumblebee colony has 50 to 200 individuals at a time. Food is stored in some of the cells, while other cells contain brood in various stages of development. Common eastern bumblebee (Bombus impatiens) buzz pollinating a Chinese beautyberry (Callicarpa cathayana) flower. 30 Arnoldia 71\/4 Buzz pollination 31 I recorded bumblebee buzz pollination on the following plants at the Arboretum. There are a number of other plants in the collections that are buzz pollinated, including Vaccinium species such as lowbush blueberry (V. angustifolium). Callicarpa cathayana Callicarpa dichotoma Callicarpa japonica Diospyros virginiana Hypericum `Hidcote' Lespedeza bicolor `Natob Strain' Rosa `Bucbi' Rubus odoratus Stewartia sinensis Chinese beautyberry Purple beautyberry Japanese beautyberry Common persimmon `Hidcote' hybrid St. John's wort `Natob Strain' shrub bushclover Carefree Beauty rose Fragrant thimbleberry Chinese stewartia The anthers of common persimmon (Diospyros virginiana), seen in the center of these flowers, are not technically poricidal but they do dehisce only partially. NANCY ROSE Fruit of purple beautyberry in autumn. 32 Arnoldia 71\/4 Buzz pollination 33 3. Why do buzz frequencies change? Could it be that different flowers require different frequencies to get maximum pollen release? Is humidity affecting the bumblebee or the flower more? 4. Why do bumblebees buzz on plants with longitudinally dehiscent anthers? Before I started collecting data, I thought that bumblebees shouldn't buzz on longitudinally dehiscent anthers since the pollen is readily accessible. But I found multiple instances of buzz pollination occurring on St. John's wort (Hypericum `Hidcote') and Carefree Beauty rose (Rosa `Bucbi'), and I even recorded some buzzing on Chinese stewartia (Stewartia sinensis). All three of those plants have longitudinally dehiscent anthers. Stephen Buchmann (1985) published similar observations and hypothesized that buzzing may increase effectiveness at collecting pollen on longitudinally dehiscent anthers, especially when the flower has a \"shaving brush\" structure (contains numerous stamens with long filaments). One suggestion for why bumblebees use buzz pollination on this type of flower is that it allows them to get pollen from many anthers at one time. With these flowers, bees gather many anthers together and hold them close to their bodies while they buzz. Though this doesn't require buzz pollination, buzzing could result in faster pollen collection than collecting from one anther at a time. Bumblebees were observed buzz pollinating the flowers of St. John's wort (Hypericum `Hidcote', seen here) even though it does not have poricidal anthers. blueberries or combing through a recipe book to find something to do with all your tomatoes, you can thank the bees. Literature Cited Buchmann, S. L. 1985. Bees use vibration to aid pollen collection from non-poricidal flowers. Journal of the Kansas Entomological Society 58: 517"},{"has_event_date":0,"type":"arnoldia","title":"2013 Weather Summary","article_sequence":4,"start_page":34,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25567","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25eab6b.jpg","volume":71,"issue_number":4,"year":2014,"series":null,"season":null,"authors":"Pfeiffer, Sue A.","article_content":"2013 Weather Summary Sue A. Pfeiffer SUE PFEIFFER A light snow in late February flocked trees along Meadow Road. JANUARY was a relatively warm and dry month. The first two-thirds of the month saw high temperatures mostly in the high 30s and 40s ( 2013 Weather 35 lating snow, which made any sort of transportation a challenge on snow-covered roads and sidewalks. Temperatures warmed during the following week, reaching above 40 36 Arnoldia 71\/4 KYLE PORT 2013 Weather 37 oppressive and uncomfortable conditions outdoors, the heat index reaching 108 Arnold Arboretum Weather Station Data 2013 Weather 39 OCTOBER was seasonably warm and very dry. The first few days of the month were warm and sunny but turned cloudy and drizzly for the following few days. Temperatures fell throughout the month, reflecting the progressing fall season. We received a light frost on the 25th, bringing the growing season to an end. The 185-daylong growing season was nine days shorter than the previous two years and the shortest growing season over the last five years. Since the first week of the month, minimal precipitation was recorded; three events produced only 0.12 inches of rain total. This lack of rain made fall leaf pick up in the Arboretum extremely easy. Supplemental watering was necessary in certain collections, but despite the lack of rain, fall color was in full swing. NOVEMBER was the fourth straight month with average low temperatures below the 30-year norm as we experienced colder than average overnight lows. Temperatures dipped below freezing (32"},{"has_event_date":0,"type":"arnoldia","title":"Simply <i>Spirea</i>","article_sequence":5,"start_page":40,"end_page":40,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25571","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25eb726.jpg","volume":71,"issue_number":4,"year":2014,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Simply Spirea Michael S. Dosmann A s I write on this mid-March day, a pile of thick, crusty snow lies on the ground while the sky continues to shower the Arboretum with more of the icy mess. It has been a long winter. But the longer days give me hope that spring is just around the corner and soon we will see the blooms of old botanical friends. One of these is Spiraea prunifolia var. simpliciflora, a delicate spirea collected as seed by John George Jack in 1905 during his trip to Korea (for more about this extraordinary plantsman, see the article starting on page 2 of this issue). Jack's original plant (accession 18283-A) still grows below a canopy of hickory trees along Valley Road. An earlier accession of the same species (accession 3138) came from the Royal Botanic Gardens, Kew, in 1887 and still grows in the Bradley Rosaceous Collection. This spirea is native to eastern Asia and grows wild throughout China, Korea, and Japan. Ernest H. Wilson had also collected the species in Korea, but that accession perished long ago. S. prunifolia var. simpliciflora is a fine-textured shrub that functions well as a single specimen, group planting, or in the mixed border. It reaches 5 to 6.5 feet (1.5 to 2 meters) tall and at least as wide. When allowed to reach its full size, the long stems grow gracefully from the center and arch up and away to create a vase shape. Cold hardy through USDA Zone 5 (average annual minimum temperature -10 to -20 "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23439","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170af6e.jpg","title":"2014-71-4","volume":71,"issue_number":4,"year":2014,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Untangling the Twisted Tale of Oriental Bittersweet","article_sequence":1,"start_page":2,"end_page":18,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25565","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25ea76d.jpg","volume":71,"issue_number":3,"year":2014,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Untangling the Twisted Tale of Oriental Bittersweet Peter Del Tredici PETER DEL TREDICI I t's amazing that the details of the introduction of one of eastern North America's worst invasive plants, Oriental bittersweet (Celastrus orbiculatus Thunb.), are essentially unknown. According to Alfred Rehder in his seminal Manual of Cultivated Trees and Shrubs (1927) the vine was introduced into cultivation from Asia in 1860, but he offered no specific details about who the responsible party was. Since then, most authors have simply taken Rehder at his word and repeated the 1860 date without question (or attribution). More recently, some botanists have cited 1879 as the date of introduction of Oriental bittersweet into North America based on an 1890 article by Charles S. Sargent, but again with only minimal details. The purpose of this article is to fill in this void in the early history of the plant, especially now that it has become such a ubiquitous--and highly destructive--member of our flora. A Brief History of Oriental Bittersweet The first species of Celastrus to be described was the American Oriental bittersweet, in yellow fall foliage, scrambles to the top of a tall eastern or climbing bittersweet (also cottonwood (Populus deltoides) in Bussey Brook Meadow at the Arnold Arboretum. called waxwork or stafftree), native to eastern North America, and named Thunberg, in his ground-breaking Flora JaponC. scandens by Linnaeus in 1753. The second ica under the name Celastrus articulatus. Some was Oriental bittersweet, C. orbiculatus, native ninety-seven years later, the Russian botanist to Japan, Korea, and China and originally pubCarl Maximowicz pointed out that this name lished in 1784 by Linnaeus's student, Carl Peter was actually a misprint of Celastrus orbicula- Oriental Bittersweet 3 tus, Thunberg's intended name, which he used in the index of Flora Japonica as well as in the original manuscript pages of the book. It took years of back and forth debate among botanists to straighten out the confusion caused by this simple typographical error, but C. orbiculatus is now universally accepted as the correct scientific name for Oriental bittersweet. In Flora Japonica Thunberg also described a second Japanese species of bittersweet, C. punctatus, with smaller, more ovate leaves than C. orbiculatus, a different pedicel (flower stalk) structure, and rough white lenticels on its stems. Shortly after this plant entered cultivation in the mid- to late 1800s, it too became engulfed in a taxonomic debate, specifically as to whether it was a \"good\" species or just a variety of orbiculatus. Alfred Rehder, writing in L. H. Bailey's massive Cyclopedia of American Horticulture (1900), officially reduced C. punctatus to a variety of C. orbiculatus, with shorter petioles and smaller, thicker, elliptic leaves. This reduction in status was widely accepted in botanical publications for many years, most notably in the English version of Jisaburo Ohwi's Flora of Japan (1965), which described variety punctatus as \"a southern phase, abundant usually near seashores, although transitional with the typical phase [orbiculatus].\" The traditional view of Oriental bittersweet taxonomy underwent a change in 1955 when Ding Hou, a freshly minted Ph.D. from Washington University in St. Louis, published his revision of the genus Celastrus in the Annals of the Missouri Botanical Garden. Hou reviewed the tortured history of Thunberg's two bittersweets and concluded they were both valid species. He also reviewed the taxonomy of the two Celastrus species described and illustrated in 1860 by Eduard von Regel, the Director of the St. Petersburg Botanical Garden: one was a \"new\" species that he christened C. crispulus, the other was Thunberg's species, C. punctatus. Writing in Plantae Wilsonianae in 1915, Alfred Rehder had expressed the opinion that both of Regel's plants belonged to the species C. orbiculatus--crispulus was a synonym and punctatus a variety--a determination that formed the basis for his citing 1860 as the date of Oriental bittersweet's introduction into cultiva- tion. Ding Hou looked at the same article and reached a very different conclusion--Regel's crispulus was synonymous with Thunberg's punctatus and his punctatus was really Thunberg's orbiculatus. According to Hou's interpretation, Rehder was right about 1860 as the date for the introduction of Celastrus orbiculatus, but wrong about which of Regel's two species was the true Oriental bittersweet. In the years following its publication, Ding Hou's revision of the genus Celastrus has stood the test of time. The current online Flora of Japan Database Project, for example, treats C. punctatus as a semi-evergreen species native to the warm-temperate or subtropical parts of the country, while the deciduous species C. orbiculatus is found in more northerly cool- and warm-temperate zones. Similarly, the English version of the Flora of China, which describes twenty-five species of Celastrus, includes both C. orbiculatus and C. punctatus. The former is widely distributed in the eastern and northeastern parts of the country, mainly north of the Yangtze River, while the latter is restricted to southeast China and Taiwan. Introduction Into Europe Eduard von Regel's 1860 Gartenflora article is significant for three reasons: 1) it is the first report of the cultivation of Oriental bittersweet outside of Asia; 2) it contains the first scientific illustrations of both Celastrus orbiculatus and C. punctatus; and 3) it unequivocally states that C. punctatus (= C. orbiculatus according to Hou) had \"only recently been imported\" into European gardens by the famous naturalist Philipp von Siebold. Siebold is an important and colorful figure in the early history of European involvement in Japan. His spectacularly illustrated Flora Japonica--co-authored with Joseph Zuccharini and published in thirty volumes between 1835 and 1870--is a botanical landmark. Siebold was a Bavarian physician who spent six years (1823 through 1829) in Japan working for the Dutch government, teaching and practicing medicine, and making a significant collection of Japanese flora and fauna. His sojourn ended when he was imprisoned for political reasons (the unauthorized possession of a strategically important Illustration of Celastrus orbiculatus and C. punctatus from Eduard von Regel's 1860 article. Oriental Bittersweet 5 map of Japan) and forced to return to Holland in 1830. He did, however, manage to leave with a boatload of herbarium specimens and living plants, which he cultivated in his garden in Leiden. Siebold managed to return to Japan in August 1859 but was forced to leave in 1862. Again, he returned to Leiden with a collection of Japanese plants that he added to the \"Jardin D'Acclimatation,\" which he had established in the 1830s (Spongberg 1990). He published a nursery catalogue for the garden in 1863 that listed an astounding 838 species and varieties of plants for sale, mainly from Japan and China. Included among the entries was \"Celastrus punctatus Thbg.\" at the price of 1 or 2 francs, presumably depending on the plant's size. Based on this catalogue listing and on Regel's article from 1860, we can now say that Siebold probably collected seeds of C. orbiculatus (which he called C. punctatus) in the fall of 1859--at the start of his second visit to Japan--and sent them to colleagues in Europe for cultivation. Siebold's 1863 nursery catalogue listing appears to be the first recorded public offering of C. orbiculatus outside of Asia. Portrait of Samuel B. Parsons from Meehan, 1887. Introduction Into North America On the other side of the Atlantic, Oriental bittersweet made its horticultural debut in the Kissena Nurseries catalogue first published in 1886 or 1887. The Kissena Nurseries were established by Samuel B. Parsons in 1871 as the successor to the earlier nursery he had established with his brother Robert in 1840 in Flushing, New York. The nursery specialized in ornamental trees and shrubs and was the first nursery in the United States to introduce Japanese maples into commerce and to propagate and distribute hardy evergreen rhododendrons (Meehan 1887). The Arnold Arboretum library has two virtually identical copies of the Kissena Nurseries \"Descriptive Catalogue of Hardy Ornamental Trees, Flowering Shrubs and Vines.\" One of them has \"1887?\" penciled on it while the other is marked \"Probably issued Spring, 1889.\" Both of the catalogues are 94 pages long and both include the identical entry for Oriental bittersweet on page 53: \"Celastrus punctatus, Japan. Leaves marked with points of white. 75 cts.\". (This reference to \"points of white\" is probably a misinterpretation of the word punctatus, which Thunberg used in reference to the prominent white lenticels on the stems.) In the Rhododendron section of the catalogue, on page 78, there is a reference to \"a recent published paper from C. M. Hovey, whose experience in this plant is well known, he states that he bought in 1884 [should read 1844], in England, a number of Rhododendrons supposed to be hardy.\" A search of the literature from this period turned up Hovey's article in the December 1885 issue of The American Garden, which makes spring 1886 the earliest possible date for the publication of the Kissena Nurseries catalogue. The listings for Celastrus from Parsons's 1887 Kissena Nursery Catalogue. 6 Arnoldia 71\/3 Oriental Bittersweet 7 Illustration of Celastrus orbiculatus by Charles Faxon from Sargent's 1890 article in Garden and Forest. Arnold Arboretum accession card for Celastrus orbiculatus accession 190 from Samuel Parsons. A check of the Arboretum's old card file system revealed that accession 190 had indeed been sent to the Arboretum by Samuel Parsons in 1879 under the name C. punctatus. In their articles, both Jack and Sargent changed the specific epithet to articulata instead of punctatus. Whether they did this because they thought the two species were synonymous or because they thought the plant was misidentified is unclear, but the latter explanation is more likely. Remarkably, the card file also revealed that seeds of \"Celas- Herbarium specimen from Arnold Arboretum accession 190-1, a plant raised from a cutting from the original plant from Parsons. Oriental Bittersweet 9 trus articulatus\" (accession 192) were received by the Arboretum on March 2, 1880, from the Agricultural College in Sapporo, Japan, less than a year after Parsons sent the Arboretum a plant of \"C. punctatus.\" Fortunately the Arboretum possesses herbarium specimens of both of these accessions, one from accession 190-1, which originated from a cutting collected on October 20, 1887, from Parsons's original plant, and the other from one of the original Sapporo plants collected on October 26, 1888. Both herbarium specimens are labeled \"articulata\" and both are in fruit, but only the Parsons specimen has leaves on it. As far as I have been able to determine, they are both Celastrus orbiculatus. Who Sent the Seeds? The unanswered question about the introduction of Oriental bittersweet into North America boils down to this: Where did Samuel Parsons get his plants? One possibility is that they came from Dr. George Rogers Hall, an American physician who lived in Japan from 1855 through 1861 and introduced many Japanese plants (including many collected by Siebold) into North America (Spongberg 1990). In March of 1862, upon his return to the United States, Hall hand-delivered a large shipment of Japanese plants and seeds to Parsons, who breathlessly described unpacking them in The Horticulturist. While there is no mention of Celastrus in the article, the door of possibility is left slightly ajar with the statement that the shipment contained \"a large number of other tree and shrub seeds.\" But this seems an unlikely source for bittersweet given that it would have necessitated a seventeen year time lag before its distribution to the Arnold Arboretum. In addition, a comprehensive article titled \"Ornamental Vines\" by Josiah Hoopes in The Horticulturist (July 1874) describes American bittersweet (Celastrus scandens) and one of Hall's notorious introductions, Japanese honeysuckle (Lonicera japonica), but makes no mention of Oriental bittersweet. The available evidence--what little there is-- suggests that Thomas Hogg, Jr. was the source of Parsons's Oriental bittersweet seeds. Hogg served as the United States marshal assigned to the Japanese Consulate from 1862 to 1869 and later as an advisor to the Japanese Customs Service from 1873 through 1875. Hogg's father, Thomas, Sr., had immigrated to New York City from London in 1821 and established one of the first nurseries in the area. When Thomas, Sr. died in 1854, his two sons, James and Thomas, Jr., took over the business. During his diplomatic appointment in Japan, Hogg used the opportunity to send a number of Japanese plants--most notably variegated hostas and Japanese irises--to the family nursery in New York as well as to other horticulturally minded individuals in the northeast (Sargent 1888, 1894; Whitehead 2011). Hogg interacted with various Japanese nurseries as well as the European botanists who were working in Japan at the time, most notably Carl Maximowicz who lived in Japan from 1860 through 1864 and collected numerous plants--including Oriental bittersweet--for the St. Petersburg Botanical Garden (Bretschneider 1898). In a letter to his brother James (published in The Horticulturist in 1863), Hogg described their relationship: \"There is a Russian Botanist (Mr. Macimovitch) now here making a collection of living COURTESY OF THE GRAY HERBARIUM OF HARVARD UNIVERSITY Portrait of Thomas Hogg, Jr. 10 Arnoldia 71\/3 Thomas Hogg, Jr.'s Plant Introductions Thomas Hogg, Jr. introduced many Japanese plants--both wild species and horticultural selections--to North America. Among his most famous are the old-fashioned variegated hostas `Decorata' and `Undulata Albomarginata', numerous Japanese maple cultivars, and the golden thread-leaved cypress (Chamaecyparis pisifera `Filifera Aurea'). Writing in the Transactions of the Massachusetts Horticultural Society for the Year 1880, Samuel B. Parsons, Jr. wrote, \"Mr. Hogg has given us possibly more new Japanese plants than any collector since the time of Robert Fortune's famous horticultural explorations.\" While I've been unable to locate a comprehensive list of Hogg's introductions, the horticultural literature of the late nineteenth century is rife with references to them. The most important sources are an article by Hogg himself in Gardener's Monthly and Horticulturist in 1879 (GMH), the 1887 Kissena Nurseries catalogue (KN), and Charles Sprague Sargent's writings in Garden and Forest (GF) from 1888 to 1897 and The Forest Flora of Japan (FFJ) in 1894. From these four references, I've compiled the following list of Hogg's woody plant introductions from Japan. No doubt persistent digging will add more species to this list in the future. Introduction years are from Rehder's Manual of Cultivated Trees and Shrubs. PLANT Veitch fir, Abies veitchii Katsura tree, Cercidiphyllum japonicum Sweet autumn clematis, Clematis terniflora Kousa dogwood, Cornus kousa Yeddo euonymus, Euonymus hamiltonianus var. sieboldianus Japanese winterberry, Ilex serrata Kobus magnolia, Magnolia kobus Japanese umbrella magnolia, Magnolia obovata Oyama magnolia, Magnolia sieboldii Japanese photinia, Photinia villosa Kudzu, Pueraria lobata Japanese hydrangea vine, Schizophragma hydrangeoides Stachyurus, Stachyurus praecox Japaneses stewartia, Stewartia pseudocamellia Sapphireberry, Symplocos paniculata Siebold viburnum, Viburnum sieboldii YEAR OF INTRODUCTION 1874 1864 or 1865 1864? 1874 1865 1866 1865 1865 circa 1865 1865 -- -- 1865 1868 1865 -- REFERENCE Sargent FFJ, p. 83 Hogg GMH 21: 53 Sargent GF 3: 621 Sargent FFJ, p. 47 Sargent FFJ, p. 26 Sargent FFJ, p. 25 Sargent FFJ, p. 10; GF 6: 65 Sargent FFJ, p. 9; GF 1: 305 Parsons KN, p. 24 Sargent GF 1: 67 Sargent GF 6: 504 Hogg GMH 21: 53 Sargent FFJ, p. 18 Sargent GF 9: 34 Sargent GF 5: 89 Sargent GF 2: 556; Parsons KN, p. 50 Sapphireberry Japaneses stewartia Kousa dogwood Color illustration of Celastrus orbiculatus from Curtis's Botanical Magazine, 1898, vol. 124 [ser. 3, vol. 54]: tab. 7599. Oriental Bittersweet 13 secured in 1897 from the Royal Gardens, Kew, granite and conglomerate were covered with England.\" Nash also noted that the painting Oriental bittersweet \"whose stems are coiled that accompanied the article \"was prepared and twisted into an intricate clump of growth, from a vine growing on some small trees in the picturesque at all season of the year.\" No doubt rear of the Museum building of the New York he was referring to plants that E. J. Palmer later Botanical Garden. It was of accidental occurreported finding on the south side of Hemlock rence there, and perhaps originated from seed Hill in his 1935 publication, Supplement to the carried by the birds from the large specimen in Spontaneous Flora of the Arnold Arboretum. the viticetum [a place where vines, especially While Wilson was an admirer of Oriental bittersweet, the Arboretum's longtime hortigrapevines, are cultivated] but a short distance culturist, Donald Wyman, was its true chamto the east\"--the very plant that had come from pion. He wrote about the plant in various Kew Gardens in 1897. So the cycle is complete: Arnold Arboretum publications in 1939, 1944, bittersweet seeds went from the wilds of Japan and 1950 as well as in a number of other horto Flushing to Boston to England and then back ticultural publications, and described it in his to New York where they began to naturalize! best-selling Shrubs and Vines for American Oriental bittersweet was a relatively rare cultivated plant towards the end of the nineteenth Gardens, published in 1949. Wyman's 1944 century, mainly confined to the properties of article was a survey of the use of rapidly growing vines in the United States, which concluded wealthy horticultural enthusiasts. With its dramatic fruit display and rampant growth, however, the plant was destined for popularity, and the staff of the Arnold Arboretum, as it had done earlier, was leading the charge. E. H. Wilson, writing in his 1925 book about the Ar nold Arboretum, America's Greatest Garden, described the plant in glowing ter ms, \"On the left ascending the Bussey Hill road, is another arresting feature. It is merely a dense tangle of Japanese Bittersweet (Celastrus articulata) but how beautiful!-- a mass of clear yellow foliage and a wild profusion of fruits with deep yellow husks cracked open, disclosing the clustered seeds clad in jackets of cinnabarred.\" Later on he notes that some of the Arboretum's boulders of A tangle of fruiting Oriental bittersweet on a stone wall in Cornwall, Connecticut. PETER DEL TREDICI 14 Arnoldia 71\/3 PETER DEL TREDICI Oriental Bittersweet Life History Celastrus orbiculatus is a high-climbing vine with stems that can grow up to 15 feet long in a single season and 60 feet long at maturity. It lacks tendrils and climbs by means of twining shoots that can eventually strangle the trunk of its host tree-- not unlike a botanical boa constrictor (Lutz 1943). Oriental bittersweet produces simple, alternately arranged leaves that are highly variable in shape--from round or egg-shaped to oblong or elliptical; they are smooth with wavy, slightly toothed margins and tips that taper to a long or short point. Bittersweet roots are shallow growing and bright orange (a good field identification characteristic) and are used as an anti-inflammatory in traditional Chinese medicine. Any piece of root that is left behind after pulling or cutting the stems will give rise, Medusalike, to numerous sucker shoots. This root-suckering capacity makes it extremely difficult to control Oriental bittersweet in landscapes where it has become established (Dwyer 1994). Oriental bittersweet produces small, greenish flowers that typically become unisexual by the developmental failure of either the male or the female organs, thus making the plant functionally dioecious (Brizicky 1964). Occasionally a plant will develop both unisexual and perfect flowers (polygamodioecious), leading to individual specimens that are functionally monoecious (Wyman 1950; Hou 1955). The inconspicuous flowers are insect pollinated (mainly by bees) and produced on lateral branches in May and June. Following pollination, female plants produce round green fruits (capsules) that become highly conspicuous in the fall when they turn yellow and then split open to reveal seeds covered with a scarlet aril. A wide variety of birds (both native and exotic) feed on the brightly colored fruits and disperse the seeds across the landscape. Seedlings are common under the trees and shrubs where birds roost at night and seeds can remain viable in the soil for several years (Dwyer 1994). Oriental bittersweet is highly adaptable and grows under a variety of light and soil conditions. Compared with the native C. scandens, the seedlings and young root sprouts of C. orbiculatus are extremely shade tolerant and can persist in the forest understory for a long time waiting for a light gap to develop (Leicht and Silander 2006). The plant is notorious for its ability to strangle and overwhelm nearby trees and shrubs and can cause serious damage in forests (Fike and Niering 1999). Oriental bittersweet was widely planted for ornamental, erosion control, and wildlife habitat purposes in the United States in the 1950s through 1970s and is now considered an invasive species throughout much of eastern North America. A recent publication from New Zealand (Williams and Timmins 2003) documented the spread of Oriental bittersweet in northern portions of that country, beginning in 1975. \"A botanical boa constrictor\"--Oriental bittersweet strangling a black locust tree. PETER DEL TREDICI Oriental bittersweet root suckers. 16 Arnoldia 71\/3 Oriental Bittersweet 17 A curtain of Oriental bittersweet foliage. Brizicky, G. E. 1964. The genera of Celastrales in the southeastern United States. Journal of the Arnold Arboretum 45: 206 18 Arnoldia 71\/3 "},{"has_event_date":0,"type":"arnoldia","title":"Magnolia virginiana: Ephemeral Courting for Millions of Years","article_sequence":2,"start_page":19,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25564","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25ea728.jpg","volume":71,"issue_number":3,"year":2014,"series":null,"season":null,"authors":"Losada, Juan M.","article_content":"Magnolia virginiana: Ephemeral Courting for Millions of Years Juan M. Losada MICHAEL DOSMANN The Arnold Arboretum's magnolia collection currently holds 157 accessions of native and non-native magnolias. A t the end of the seventeenth century, Henry Compton, the Bishop of London and a man known for his passionate love of gardening, sent the Reverend John Banister on a missionary trip to the New World. Banister arrived in Virginia in 1678 and, in addition to his clerical work, collected many new plant species for Bishop Compton. Among these was a tree species never before seen in Europe, specimens of which were planted and flourished near Fulham Palace, the Bishop's residence. After observing these specimens, botanist Philip Miller recorded the first written reference to this species in his book, The Gardeners Dictionary. Miller was not only the chief gardener of the Chelsea Physic botanic garden, the second oldest in Britain, but also a plant collector and conservationist who cultivated many exotic species. Interestingly, the garden was visited by Linnaeus during his trip to England in 1736. Miller was influenced by the new system of classification that Linnaeus proposed, to the extent that he organized the garden following the Linnaean system. In 1753, Linnaeus included for the first time in his world renowned work Species Plantarum the specimens that Miller observed at Fulham Palace, with the name Magnolia virginiana. 20 Arnoldia 71\/3 Magnolia virginiana 21 nated terrestrial ecosystems. Both gymnosperms and angiosperms are seed plants, and seeds are the product of fertilized ovules. Herein lies the main difference between both plant groups: while gymnosperms have their ovules exposed or \"naked,\" flowering plants developed maternal tissues to shelter their ovules. Despite the more complicated new arrangement of the ovules, flowers increased the efficiency of sexual reproduction and opened up many new opportunities for coevolutionary relationships between flowering plants and insect pollinators. Flowers became key evolutionary innovations, opening a door for innumerable new reproductive strategies that can be seen throughout the great diversity of flowering plants. Sexual Reproduction and Flower Receptivity In 1694, Rudolf Jakob Camerarius published his discovery that plants undergo sexual reproduction. Flowers are the reproductive parts of angiosperms, performing two main functions: they act as a showy display to attract pollinators, and they bear the germ lineages (gametes). The germ lineages are housed inside of a number of tissues specialized for either dispersal (for the male gametes) or protection (for the female gametes). The contact of both male and female gametes in most flowering plants involves the transfer of pollen between individuals, which is a task often carried out by insect pollinators. The first major studies on plant pollination were done by K 22 Arnoldia 71\/3 Magnolia virginiana 23 within a flower--self pollination (the equivalent of marrying a very close relative). But in Magnolia virginiana (and in other Magnolia species), a temporal separation of the activities of the male and female parts of individual flowers acts to diminish the possibility of inbreeding. The temporal separation of both sexes is manifested as a protogynous flowering cycle (proto = first, gynoecium = female parts, or \"ladies first\"), and is delimited by floral movements. As a result, the female phase precludes the male phase and they do not overlap, thus creating a two day flowering cycle. Flowers open the first day at dusk (opening takes around 20 minutes and can be observed by just staring patiently at the right flower) as females with wet, sticky stigmas that receive pollen grains, and then close when night falls. They remain closed until the evening of the following day, when flowers reopen in the male phase, at which point stamens shed pollen. During the stage in which the flower remains closed, the flowers generate heat in order to give Female phase of Magnolia flowers. (A) A Magnolia flower shows multiple gynoecia at the first flower opening. (B) Detailed view of the hooked stigmas that shelter to their main pollinator, have bright, sticky surfaces ready to receive pollen grains. (C) Scanning electron beetles. The ability for flowers micrograph of the stigma surface in Magnolia virginiana, showing the fingerlike to produce heat is common to all cells (papillae) that form an intricate network for pollen grain gathering. magnolias (and other members of actions, recorded with time-lapse photography the family), and so is thought to be an ancestral character for the lineage. Other pollinators, under controlled conditions The resulting video such as bees, have been observed to act as polis available online: linators for these plants, but little is known http:\/\/www.youtube.com\/ about how effectively they transfer pollen from watch?v=Ja3GJyJ98uI flower to flower. A few studies in the reproductive biology of The timing of flower movements affects the genus Magnolia suggested that the period reproductive performance and points to the of female receptivity was connected to these importance of a rhythm. This rhythm could flower movements, but exact timing was be associated with pollinator behavior, in our unknown. Our investigations in the Arborecase mainly bees and bumble bees, and possibly tum with controlled pollinations in the laborabeetles. Our research project with M. virgintory confirmed those suggestions, and showed iana at the Arnold Arboretum started with the that stigmatic receptivity is remarkably short. observation of this cycle and pollinator inter- 24 Arnoldia 71\/3 Magnolia virginiana 25 composed of two organic units: small amino acid backbones, and large sugar moieties where the functional capacity resides), which have numerous functions in plants such as acting as mediators in cell-to-cell communication, were secreted towards the apple stigma surface precisely at the time of receptivity. Furthermore, these glycoproteins are known to control plant cell elongation processes, and could be involved in pollen tube elongation. Their conspicuous presence in female tissues of apple flowers prompted us to wonder whether ancient lineages of angiosperms (flowering plants) would use similar molecular mechanisms. Microscopy evaluation of Magnolia virginiana stigmas showed that the nutrient movements in stigmatic tissues followed a precisely defined cycle, and that the secretory products on the stigma surface were mainly saccharides (short chains of sugars, based on the binding of individual units such as glucose or fructose). Furthermore, by using antibodies (immunolocalization) specific for the glycoproteins that were also present in apple stigmas, we detected these molecules during the short period of stigmatic receptivity in M. virginiana. This suggests that in M. virginiana, as in apple, specific glycoproteins mark the short time frame that flowers are able to allow pollen grain germination on the stigmatic surface. This work showed for the first time in a member of the Magnoliaceae that maternal tissues bear glycoproteins during pollen reception, and hinted at their involvement in pollen tube elongation towards the ovules. Combined, all this data offers new perspectives on how different flowering plants control the production of offspring. The presence of common nutritive factors secreted from the female tissues at times of pollen reception in very distantly related species points to a possible conserved mechanism across all angiosperms. But also, it sheds light on the molecular crosstalk during initial stages of male The Evolution of Pollen Receiving Structures in Seed Plants SHOWN HERE are illustrations of longitudinal median sections of different maternal tissues receiv- ing pollen grains in seed plants. The associated cladogram shows the estimated time of emergence for general seed plant lineages (mya=millions of years ago). The earliest group shown is the gymnosperms, which arose around 290 million years ago, and are characterized by naked ovules that have a liquid secretion at their ovule tips (the pollination droplet) directly catching pollen grains. Those pollen grains germinate following contact with ovule tissues. In contrast, angiosperms evolved around 243 million years ago, and most basal flowering plants had already developed maternal tissues surrounding their ovules. Among them, the apical part (the stigma) establishes the first contact between maternal tissues and paternal pollen grains. In the basal angiosperm lineages (Amborellales, Nymphaeales, and Austrobaileyales), the stigmas produce a copious secretion at their surface for pollen reception. More evolved but still relatively early divergent angiosperms show large stigmatic surfaces and a wet appearance, but lack a copious secretion. Pollen grains can develop different pollen tube lengths depending on the area of the stigma where they are deposited. Finally, in most evolved angiosperms (in a broad sense), stigmas tend to reduce their area, whereas larger styles developed, and a specialized central transmitting tissue is the arena for pollen tube elongation towards the ovules. These illustrations emphasize the importance of the stigma during the first male Magnolia virginiana 27 when they included this species in their palace gardens. However, they missed the equally remarkable story behind what was happening within those flowers: the impressive coordination of floral movements and molecular interactions that created the ephemeral female phase, a short time for a courtship repeated every blooming period for millions of years. Acknowledgements I am very grateful to Ned Friedman and Becky Povilus for valuable comments and their contribution to the fluency of this manuscript. References Camerarius, R. J. 1694. De sexu plantarum epistola. Tubingen. Darwin, C. R. 1862. Fertilisation of orchids. London: John Murray. Darwin, C. R. 1876. The effects of cross and self fertilisation in the vegetable kingdom. London: John Murray. Friedman, W. E. 2006. Embryological evidence for developmental lability during early angiosperm evolution. Nature 44: 337"},{"has_event_date":0,"type":"arnoldia","title":"Wish You Were Here","article_sequence":3,"start_page":28,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25566","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25eab26.jpg","volume":71,"issue_number":3,"year":2014,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"Wish You Were Here Nancy Rose L ong before email, Twitter, or Instagram, postcards were the medium of choice for sending brief messages and colorful images to friends and family. Simple cards, mostly used for advertising, were first introduced in the United States in the 1860s but were not especially popular since they required the same postage as letters. In 1873, the United States Postal Service (USPS) introduced official \"postal cards,\" plain cards with a printed stamp. The postage cost was one cent, half the rate for letters. However, non-USPS cards still required the full two-cent rate. The rise of souvenir postcards can be traced to the 1893 World's Columbian Exposition in Chicago, where vendors offered USPS postal cards with the addition of full color images of Exposition sights printed on the front. Finally, in 1898, an Act of Congress allowed privately printed postcards to mail at the same rate as USPS cards. Over the following decade the popularity of postcards soared, starting to decline only with trade and tariff issues prior to and during World War I (most postcards of the time were printed in Germany) and the increasing prevalence of telephones in the 1920s. Given the beauty of the Arnold Arboretum it's not surprising that it has been featured on many postcards over the years. The Arboretum archives hold a folder full of these historical postcards, some of which are presented below. Linen style postcards were introduced in 1931. They are notable for the fabric-like texture embossed on the paper and their crisp, bright colors. This postcard of the Arboretum's Hunnewell Building (then simply called the Administration Building) was likely produced in the 1930s but was clearly based on a 1921 black-and white photograph made by Alfred Rehder (the vine coverage on the building matches precisely!). Postcards 29 One hundred forty characters or less? Prior to 1907, postal regulations allowed only the mailing address to be written on the stamped side of cards. On many souvenir cards, whose main appeal was the colorful image on the front, this left only a narrow strip at the bottom for a personal message. This may have inspired concise composition (the upper card reads \"With kind regards, hope you are all well, from your friend Jemima Cook\") or very tiny lettering. The lower card shows the pre-1907 admonition against writing messages on the stamped side but by 1910, the year it was postmarked, the sender's message--\"Dear Grandpa, Mamma send[s] her love and hopes you are well\"--was perfectly legal. 30 Arnoldia 71\/3 Postcards 31 In living color The original postcard boom was in part related to the development and proliferation of chromolithography in the latter nineteenth century. Using multiple lithographic stones or plates to apply layers of color, this printing process greatly increased the availability of high quality but affordable color prints. The best quality chromolithograph postcards were printed in Europe, primarily Germany. Hand colored images could be printed in all their glory on postcards, though printers could be variable in color quality. Among the Arboretum cards, some show fairly natural colors while others have little resemblance to the actual landscape. These two cards (facing page) show the same view from within the Arboretum's lilac collection on Bussey Hill, looking east toward the ponds and Forest Hills gate. The top card was made in Germany circa 1906 32 Arnoldia 71\/3 Postcards 33 Where in the Arb? Hundreds of thousands of postcards featuring everything from local taverns to the Grand Canyon have been printed over the past 100-plus years. Thanks to long-standing appreciation of postcards as collectors' items, a surprising number of these bits of paper have been preserved. Postcards have come to be recognized as valuable research materials for historians of architecture, landscapes, and other natural and man-made features, including places like the Arboretum. It's particularly interesting to see the same view over the years: Seen here, a beautiful circa 1907 34 Arnoldia 71\/3 Postcards 35 We regret the error Poor spelling and misinformation did not originate with the internet--even the early postcard era had its share of errors. At left, a card incorrectly names the Arboretum's Bussey Hill as \"Buzzy Hill,\" while the rather unappealing card below labeled \"Scene in Arnold Arboretum\" appears to be the entrance to nearby Franklin Park instead. Suggested Reading Prochaska, D. and J. Mendelson, eds. 2010. Postcards: Ephemeral Histories of Modernity. University Park: Pennsylvania State University Press. Stevens, N. ed. 1995. Postcards in the Library: Invaluable Visual Resources. New York: Haworth Press. Willoughby, M. 1992. A History of Postcards: A Pictorial Record from the Turn of the Century to the Present Day. London: Studio Editions. Nancy Rose is the editor of Arnoldia. "},{"has_event_date":0,"type":"arnoldia","title":"Chimonanthus praecox: A Redolence of China","article_sequence":4,"start_page":36,"end_page":37,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25563","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d25ea36f.jpg","volume":71,"issue_number":3,"year":2014,"series":null,"season":null,"authors":"Yih, David","article_content":"Chimonanthus praecox: A Redolence of China David Yih O n a raw, wintery day last February, I traveled from Connecticut to visit the Arnold Arboretum, impelled by curiosity. In 1977, my father, at the behest of the poet Donald Hall, had written a series of vignettes for The Ohio Review recalling the China he had left more than thirty years earlier. Among these was a nostalgic essay in which he sought to convey a feeling for Chinese esthetics as exemplified by Chimonanthus praecox, known in China as la mei. Its English common name, wintersweet, encapsulates two notable features of the plant: its membership in that small fraternity of temperate shrubs that bloom in winter and the remarkable fragrance of its flowers. I had recently learned that a specimen grew at the Arboretum and wanted to experience this fragrance for myself. No account of wintersweet fails to mention the scent of its blossoms. But, as my father's essay points out, the resources of the English language are scarcely adequate to describe the smell of flowers. His attempt begins by contrasting wintersweet with gardenia, orange, and locust, whose scents \"have something sensual in them that makes you feel restless, as if there were something missing in your life.\" The wintersweet's fragrance is something \"entirely different, because it is ethereal, spiritual, otherworldly.\" This distinctive scent had set off a Proustian tumult of memories when my father happened to visit a botanical garden while living in Geneva, in 1964: \"As I wandered about I suddenly smelled a remembered fragrance ... In the tepid sun and the breeze, I suddenly recalled my grandfather's house with its two wintersweet trees, my middle school in Soochow with its ancient garden, and the hills of the Chia-ling River. My mind was drunk with memories of people who had gone out of my life and of sceneries I should in all likelihood never see again.\" Chimonanthus belongs to Calycanthaceae, a small family whose members are found primarily in East Asia and North America. Endemic to montane forests in China, Chi- monanthus praecox has been cultivated for over a thousand years. A great number of cultivated varieties exist in China, where it is grown as a garden shrub, a potted plant, and for flower arrangements. When the Sung dynasty poet Huang T'ing-chien composed a poem in praise of la mei, the plant attained instant fame and popularity in the capital, Kaifeng. Fan Chengda included it in his botanical treatise, Fancun meipu (Fan-Village plum register), circa 1186. According to the custom of associating a plant with each month of the lunar calendar, la mei is the flower of the twelfth month; its blooming thus coincides with the Chinese New Year. The Arnold Arboretum's lone specimen (accession 236-98) was grown from seeds received from a botanical garden in Belgium. Wintersweet is marginally cold hardy in USDA Zone 6 (average annual minimum temperature 0 to -10 "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23438","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170af28.jpg","title":"2014-71-3","volume":71,"issue_number":3,"year":2014,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Strange Range of Seaside Alder","article_sequence":1,"start_page":2,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25562","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24e896b.jpg","volume":71,"issue_number":2,"year":2013,"series":null,"season":null,"authors":"Jones, J. Matthew","article_content":"The Strange Range of Seaside Alder J. Matthew Jones P lant species exhibit considerable variation in their native ranges. Some species, such as the North American white oak (Quercus alba), are abundant and have a continuous range across half the continent. Other species, like the dawn redwood (Metasequoia glyptostroboides) in China, are rare and natively restricted to a small location in the wild. In addition to being rare, some species exhibit disjunct, or widely separated, populations. Disjunct populations can arise either by unusual dispersal events, or can be remnants of a formerly more widespread range that has since retracted. Disjunct populations of a given species are often highly isolated from one another, which can reduce the likelihood of pollen and seed dispersal (and thus gene flow) among populations. Consequently, isolated populations are more prone to inbreeding and random selection, which, over time reduces genetic diversity and a population's ability to adapt to environmental change. Therefore, studying disjunct Delaware A. maritima ssp. maritima Maryland Oklahoma A. maritima ssp. oklahomensis Georgia A. maritima ssp. georgiensis Seaside alder (Alnus maritima) comprises three small, widely disjunct populations in Maryland, Georgia, and Oklahoma. Seaside Alder 3 Introducing Seaside Alder: The Alder That Does Not Grow by the Seaside Seaside alder is among the most disjunctly distributed tree species in North America, with major populations so widely separated that they are recognized as distinct subspecies. Alnus maritima subsp. maritima occurs on the Delmarva Peninsula, specifically Foliage and catkins of Alnus maritima. along tributaries and streams of the Naticoke, Pocomoke, and Wicomico Alnus maritima and other members of the Rivers of Maryland's Eastern Shore, as well birch family (Betulaceae, 230 species worldas along rivers, ponds, and dammed creeks in wide) are recognized by their distinctive floral Delaware. Alnus maritima subsp. oklahomencatkins and conelike infructescences. Stamisis is restricted to four populations on the Blue nate (male) inflorescences are elongate catkins River and two of its tributaries in south central that hang loosely at the tips of branches, typical Oklahoma, while A. maritima subsp. georgienof trees that disperse pollen by wind. The tiny, sis occurs along opposite sides of a single pond apetalous, pistillate (female) flowers sit above in Bartow County, Georgia. What historical, woody, scalelike leaves (bracts), arranged as a ecological, and physiological factors caused this spike in the leaf axils. At maturity, the woody disjunct distribution? How genetically diverse bracts on the \"cones\" spread apart to release the are each of the regional subspecies and how water-dispersed seeds. Unlike birches, which might this affect their long-term viability? have deciduous three-lobed bracts, alders' fiveAlnus maritima is a large shrub or small tree lobed woody bracts persist well into the folthat tends to grow in spreading, multi-stemmed lowing season, yielding an attractive display of clumps in wet soils along banks of ponds, inch-long spent \"cones\" in winter. streams, and even fast moving rivers. Seaside Eight species of alder occur in North America. alder's roots can tolerate partial submergence in Flower-to-seed development of most of these water and, similar to legumes, have a symbiotic species requires a full year. Floral buds initiate relationship with bacteria that assimilate atmoin summer, go dormant over winter, open for spheric nitrogen. Despite its name, A. maritima pollination the following spring, and disperse doesn't occur on the seaside and is intolerant their seeds in late summer and autumn. Alnus of saline soils. Individual trunks are usually 10 maritima, however, initiates floral developto 13 centimeters (4 to 5 inches) in diameter, ment in spring, opens for pollination in late 5 to 7 meters (16 to 23 feet) tall, and covered summer to early autumn, and releases seeds in in smooth, light gray bark. The simple, alterlate autumn or early winter of the same year. nate leaves are narrowly elliptical to obovate in This phenological sequence (supported by more shape, with a lustrous surface, leathery texture, recent genetic data) delineates the alder suband singly serrated margins. genus Clethropsis, a clade that A. maritima ROBERT H. MOHLENBROOK, USDA-NRCS PLANTS DATABASE populations is important for understanding how habitat fragmentation affects biodiversity and species' long-term viability, as well as for helping to assess conservation strategies that maintain genetic diversity. Seaside Alder 5 shares with the Himalayan alder (A. nitida) and the Formosan alder (A. formosana). This means that seaside alder's nearest extant relatives are in southern Asia (Chen and Li 2004). Investigating the Causes and Consequences of the Disjunction Schrader and Graves (2002) classified the three regional populations of seaside alder as three subspecies. Subspecies are a taxonomic ranking used to distinguish geographically isolated populations of species that often have unique morphological features and specialized adaptations to local environments. Alnus maritima subsp. oklahomensis is the most distinctive of the three subspecies, with narrower leaves and larger trunks and canopies compared to the Delmarva (A. maritima subsp. maritima) and Georgia (A. maritima subsp. georgiensis) subspecies. A. maritima subsp. oklahomensis also occurs in more phosphorus deficient and alkaline soils than the other subspecies. Alnus maritima subsp. maritima was the first population recognized in the early nineteenth century. Seaside alder's strange distribution was first noted when Elihu Hall discovered populations along the Blue River in the Indian Territory during his exploration of the region in 1872. Was this the result of a dispersal event or the remnant of a formerly broader range? John Furlow (1979), in his comprehensive monograph of American alder species, proposed that the Oklahoma population was a relict of range retraction, noting the fossil evidence of related species further west. Virginia Stibolt's (1981) study of seaside alder distributions suggested the Oklahoma population might be the result of dispersal by Native Americans, as the Delaware Indians were forcibly relocated to the Indian Territory several decades before Hall's discovery. However, Stibolt herself expressed skepticism about this hypothesis, and there is no ethnobotanical or historical reasoning for Native Americans to have intentionally or inadvertently dispersed seaside alder seeds. The discovery of the Georgia population in 1997 further undermined the dispersal hypothesis. Alnus maritima subsp. oklahomensis has narrower leaves and larger trunk and canopy size than the other two subspecies of seaside alder. Population Genetics as a Research Tool Population genetics, the study of genetic diversity within and among populations of organisms, has further clarified the nature of seaside alder's present disjunct distribution. Additionally, using population genetics techniques can help conservation biologists assess the genetic health of the population. Namely, how diverse and genetically distinct are the existing populations, are those populations highly inbred, and how much do populations contribute to overall species diversity? To measure population diversity and health, population geneticists compare frequencies of alleles (variants of genes) and frequencies of heterozygosity (possessing two alleles of a gene Facing page. Charles Edward Faxon's illustration of Alnus maritima in Charles Sprague Sargent's Flora of North America, 1890. In the entry for this species Sargent wrote, \"Its brilliant foliage and its bright golden staminate aments [catkins], hanging in September from the ends of the slender leafy branches, make it at that season of the year an attractive ornament for parks and gardens.\" J. PHIL GIBSON 6 Arnoldia 71\/2 Seaside Alder 7 al. (2008) suggested the alders' perennial, multistemmed, clonal growth habit and ability to sprout new shoots after disturbance and damage promote greater gene flow (movement of pollen or dispersal of seeds) and maintenance of genetic variation within each region. Additionally, genetic variation within regions might be maintained by extensive gene flow among populations within regions. In other words, although the vast distances among seaside alder in Delmarva, Georgia, and Oklahoma severely limit the exchange of pollen or seed, more localized gene flow seems to occur among networks of populations within regions. However, allozymes lack sufficient variation (or resolution) to detect differences among populations within regions, or among subpopulations within those populations. To test this hypothesis, more variable genetic markers are required. Simple Sequence Repeat Microsatellites: High Definition Population Genetics Microsatellites (also known as Simple Sequence Repeats, or SSRs) are regions of repeated sequences of DNA (e.g., GAGAGA) that do not code for proteins or enzymes. Such genes may serve some regulatory functions, but are often colloquially considered to be \"junk\" DNA. Their repeated sequence motifs lend them to a type of regular, neutral mutation that makes microsatellites particularly useful molecular marker for population genetic studies. Although more expensive and time consuming to develop and analyze, microsatellites offer much greater variability than allozymes and allow for finerscale genetic analyses. My own thesis work (Jones and Gibson 2011, 2012) used microsatellite markers to corroborate population genetic characteristics observed in J. PHIL GIBSON A colony of Alnus maritima subsp. oklahomensis grows along the Blue River in Oklahoma. 8 Arnoldia 71\/2 Seaside Alder 9 reduced the uniform dispersal of pollen clouds, a common phenomenon affecting the mating systems of wind-pollinated species. However, while these barriers tended to favor siring by fewer local pollen parents, we were able to identify potential individual pollen parents located some distance from the mother tree. In Oklahoma, some offspring collected from the main Blue River population were potentially sired by pollen parent trees located 5 to 7 kilometers (3 to 4 miles) away in nearby creeks. In Georgia, identified pollen parents were located on both sides of the pond. This means that gene flow via movement of pollen is possible among populations within each subspecies. But How Did It Get There? Both the allozyme and microsatellite genetic data strongly suggested the three regional seaside alder subspecies are the result of range retraction, not dispersal. Each population showed comparable levels of genetic diversity and each contained alleles unique to the region. What then, made seaside alder's range retract so much, and why was it so previously widespread? Why is a species like Alnus serrulata widespread, but Alnus maritima rare? Alnus maritima and Alnus serrulata share many ecological features well suited for colonizJOSEPH O'BRIEN, USDA FOREST SERVICE, BUGWOOD.ORG ing disturbed riparian sites, including tolerance of wet soils, the ability to fix nitrogen, and a robust multi-stemmed growth habit. However, A. maritima has been observed to only inhabit areas with full sun, whereas A. serrulata can inhabit both sunny and shady niches. Schrader et al. (2006) investigated the photosynthetic and growth characteristics of both alder species and found seaside alder to have lower chlorophyll concentrations, greater seedling growth rates in full sun, and lower seedling survivability in shade compared to Alnus serrulata. Therefore, while seaside alder would be more successful on relatively open, early-succession habitats, as the ecosystem matures, later succession species establish and develop canopies that shade seaside alder and reduce its competitiveness. It would eventually succumb to other shade tolerant species, including hazel alder. Schrader et al. (2006) proposed that the late Pleistocene, a time of significant climatic instability characterized by cycles of glacial advancement and retreat and changes in the locations of drainage basins, would have provided many opportunities for seaside alder to colonize and establish a broad range across the continent. Once the climate stabilized, however, those disturbance events stopped, and with it seaside alder's chance to thrive and spread. Stability brought on ecological succession, and seaside alder was gradually outcompeted by other species, except in its present range. Alders bear persistent, conelike infructescences. Those of the widely adaptable hazel alder (Alnus serrulata) are seen here. Conservation Implications The distinct genetic identity of each subspecies and the gene flow among populations within each region means that it is important to protect all populations to conserve the genetic diversity and long-term viability of the species. The Georgia and Oklahoma subspecies are now both classified as \"critically imperiled,\" the most threatened conservation status according to state conservation agencies. Recent 10 Arnoldia 71\/2 COURTESY OF STANLEY A. RICE Seaside Alder 11 A five-year-old hedge of Alnus maritima `September Sun' shows off glossy foliage and golden fall-blooming staminate catkins. Gibson, J. P., S. A. Rice, and C. M. Stucke. 2008. Comparison of population genetic diversity between a rare, narrowly distributed species and a common, widespread species of Alnus (Betulaceae). American Journal of Botany 95: 588"},{"has_event_date":0,"type":"arnoldia","title":"Alfred Rehder: His German Roots","article_sequence":2,"start_page":12,"end_page":17,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25558","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24e816f.jpg","volume":71,"issue_number":2,"year":2013,"series":null,"season":null,"authors":"Grimmer, Arnd Rudiger","article_content":"Alfred Rehder: His German Roots Dr. Arnd R Alfred Rehder 13 A.WINKLER, WWW.LUFTBILDER-WINKLER.DE Waldenburg castle, where Alfred Rehder was born, as it appears today. mate son of Heinrich, Duke of Anhalt-K 14 Arnoldia 71\/2 Alfred Rehder 15 Rehder's Years of Travel In March 1884, the apprenticeship was finished. Fourteen years of traveling then began for the young gardener, and during this period Rehder made himself familiar with all aspects of gardening work. First, he went to Berlin at the Botanical Garden of the Friedrich-Wilhelm-University and used the opportunity to gain new knowledge about taxonomy and field research. As first proof of his growing taxonomic skills, he gave a lecture at the 1886 meeting of the Association of Natural History in Zwickau. The 23-yearold spoke about beardmosses, and showed convincingly that the esteemed explorer Henry M. Stanley had made errors in the determination of this plant species (Anonymous 1886). After an interlude in 1886 working with a flower grower in Frankfurt, he went to Muskau and worked for a year with Gustav Schrefeld (1831 Brockengarten Alfred Rehder was instrumental in the creation of the Brockengarten, a botanical garden designed for the study of alpine plants. After years of neglect in the 1970s and 1980s, the garden was reestablished in the 1990s. Today, alpine plants from many regions can be seen in the garden, with a weather station and telecommunications tower arising at the mountain's peak. Seen here, sky blue Gentiana ternifolia from China and Calceolaria uniflora from the Patagonia region in South America. PHOTOS COURTESY OF DR. GUNTER KARSTE, NATIONAL PARK HARZ However, the collaboration with his supervisor, Albert Peter (1853 Alfred Rehder 17 ARCHIVES OF THE ARNOLD ARBORETUM A New Beginning in America In the 1880s, grape phylloxera (an aphid-like insect that can damage or kill grapevine roots) spread in Germany and threatened to destroy the wine producing industry. American grape (Vitis) species were found to be resistant to phylloxera, so knowledge of these plants was indispensable for the survival of European wine production. In 1898 Rehder received from the German government a mandate to examine the Vitis species on the East Coast, while at the same time reporting about American horticulture for his journal. As a destination, he chose the Arnold Arboretum in Boston, which was part of Harvard University and, despite being in existence for only 26 years, was regarded as the center of American dendrology. His initial reception in Boston was far from friendly, as the over-cautious port authorities had him summarily detained. Through the mediation of the founder and director of the Arnold Arboretum, Charles S. Sargent (1841"},{"has_event_date":0,"type":"arnoldia","title":"Remembering Alfred Rehder","article_sequence":3,"start_page":18,"end_page":24,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25560","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24e856d.jpg","volume":71,"issue_number":2,"year":2013,"series":null,"season":null,"authors":"Pearson, Lisa","article_content":"Remembering Alfred Rehder Lisa Pearson A lfred Rehder sailed for America on the Cunard steamship Cephalonia in the spring of 1898. He had a small stipend from the gardening journal he worked for, M Alfred Rehder next to a rather sprawling specimen of Magnolia stellata in front of the Arboretum's Hunnewell Building. Photograph by R.W. Curtis, June 8, 1922. seeds, cuttings, bulbs, and roots of over 1,000 of the College of Agriculture at the University species, some of them completely new, and of California, Berkeley, and later Director of the all of which needed identification and classiArnold Arboretum) Rehder laments, \"Like you fication. The documentation of this labor by I am getting `snowed under' with material coming in all the time.\" With the death of Charles Rehder and Wilson, as well as their colleagues Faxon in 1918, Rehder was appointed curator George Russell Shaw and Camillo Schneider, of the herbarium and during his 22-year tenwas the three-volume Plantae Wilsonianae: An ure he increased the holdings by over 300,000 Enumeration of the Woody Plants Collected mounted specimens. Many of those sheets were in Western China for the Arnold Arboretum of Harvard University During the Years 1907, 1908, and 1910 by E. H. Wilson. Rehder and Wilson would also collaborate on A Monograph on Azaleas, published in 1921, as well as a number of articles in the Journal of the Arnold Arboretum. In addition to their professional collaboration, they and their families were also genuine friends. In his letters to Rehder from Japan, Wilson often sends greetings from his wife Ellen to Rehder's wife Anneleise, and in a letter to Wilson from Breslau in 1930, Rehder familiarly tells about his having some \"good Rhine wine.\" Wilson's collecting trips to Japan in 1914, and Japan, Korea, and Formosa (Taiwan) in 1917 Plant Hardiness Zone Maps TODAY, plant hardiness zone information is found on everything from catalogs to plant labels in nursery pots, but less than 100 years ago there was no national map of hardiness zones for the United States. It was not until the publication of Alfred Rehder's Manual of Cultivated Trees and Shrubs Hardy in North America in 1927 that one became available, its publication predating the first USDA hardiness map by more than 30 years. This early map was divided into eight zones and did not include Florida, the southern portions of Texas and Louisiana, and it only took in the parts of Canada contiguous with the northern border of the United States. For the second edition of his Manual, Rehder used a modified version of a zone map published in 1936 by Donald Wyman that included more of Canada. ARCHIVES OF THE ARNOLD ARBORETUM The plant hardiness zone map, with colored pencil added by Rehder, in his personal copy of his 1927 Manual of Cultivated Trees and Shrubs Hardy in North America. the product of Arboretum-sponsored expeditions such as those by Wilson and Rock, but through his network of colleagues at other institutions, Rehder also actively collected duplicate sets of their sheets to fill gaps in the Arboretum herbarium holdings of the flora of important regions. Rehder as Writer and Editor In 1919, Rehder took over the behind-the-scenes management of a new institutional periodical, the Journal of the Arnold Arboretum. He had lobbied for its creation to fill the gap left by the demise of Garden and Forest in 1897 and to provide a quarterly forum for articles more technical and lengthy than could be accommodated by the Bulletin of Popular Information. In his preface to volume one, issue one, Sargent summed up its purpose, \"In its pages will appear notes on trees and shrubs with descriptions of new species and their relationships, letters from correspondents, and notes on the vegetation of countries visited by officers and agents of the Arboretum.\" While it was Sargent's name which appeared as editor at that time, the bulk of the production work, preliminary editing, and the authorship of many articles for the Journal fell on Rehder's shoulders. In 1926, Rehder Alfred Rehder 21 COURTESY OF WWW.SEQUIMRAREPLANTS.COM More than 60 plant taxa have been named in honor of Alfred Rehder. Seen here is Clematis rehderiana, a handsome clematis native to China and Nepal. became joint editor to assist an increasingly frail Charles Sargent. With volume eight the next year, Rehder and Wilson assumed joint editorial control. Wilson's untimely death in an automobile accident in October 1930 caused more reorganization and Rehder assumed the role of senior editor with Joseph Horace Faull and Karl Sax as associate editors. In the following year Clarence Kobuski, Rehder's assistant in the herbarium, took over for Sax as joint editor and continued in this role until Rehder's retirement in 1940. At that time, A. C. Smith became editor as well as curator of the herbarium, and a ten-member editorial board was formed that included Alfred Rehder, who continued in this capacity until 1948, just a year before his death. Charles Sprague Sargent, who had guided the Arnold Arboretum since its founding in 1872, died in March 1927 after several years of declining health. The institution was on shaky fiscal ground, forcing acting director Ernest Wilson to cut Rehder had blank pages bound into his personal copy of the Manual of Cultivated Trees costs wherever he could. He and Shrubs Hardy in North America so that he could write notes and corrections. trimmed the staff in Jamaica Plain and abroad, even curtailing Joseph Rock's plant collecting in western China. But in the midst of the turmoil, publication of Rehder's Manual of Cultivated Trees and Shrubs Hardy in North America provided a bright spot for the institution in that momentous year. The Manual presented \"a systematic and descriptive enumeration of the cultivated trees and shrubs hardy in North America,\" and facilitated \"their identification by means of analytical keys.\" It immediately became the go-to book for botanists and horticulturists alike. The book proved so useful and popular that it went to a second printing. In 1940, a completely revised second edition was issued, which took in new species and applied changes to rules and nomenclature adopted in the 1930 and 1935 International Botanical Congresses. Multiple reprintings of this second edition of the Manual have been made over the years, including a paperback version as recently as 2001. In the foreword to a commemorative 1986 reprinting of the Manual, botanist Theodore Dudley noted that Rehder \"possessed an insatiable curiosity and outstanding originality, demanded of himself the very highest standards, and was dedicated to the systematics and biology of all woody plants.\" ARCHIVES OF THE ARNOLD ARBORETUM 22 Arnoldia 71\/2 Alfred Rehder 23 Finding the Dwarf Alberta Spruce IN 1904, Alfred Rehder and John George Jack, Professor of Dendrology at the Arnold Arboretum, went on a plant collecting expedition to the Canadian Rockies. While waiting for a train near Lake Laggan, Alberta, they decided to take a walk. On their stroll they came upon a remarkable dwarf form of the local variety of white spruce (Picea glauca var. albertiana). Seedlings were sent back and the one surviving plant grew into a fine specimen. With its dense, almost perfectly conical growth, extreme cold hardiness, and ease of propagation, dwarf Alberta spruce (P. glauca var. albertiana f. conica) was soon adopted by the nursery trade and is now ubiquitous in gardens and container plantings. Two accessions (11586 and 182-2005) of this popular garden plant may be found in the Arboretum's Conifer Collection and the Leventritt Shrub and Vine Garden. ARCHIVES OF THE ARNOLD ARBORETUM NANCY ROSE A stand of Picea glauca var. albertiana photographed by Alfred Rehder in August 1904 near Banff, Alberta, and a potted specimen of the now widely-grown dwarf Alberta spruce. 24 Arnoldia 71\/2 "},{"has_event_date":0,"type":"arnoldia","title":"Dispersal","article_sequence":4,"start_page":25,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25559","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24e8528.jpg","volume":71,"issue_number":2,"year":2013,"series":null,"season":null,"authors":"Laurent, Anna","article_content":"Dispersal Anna Laurent Editor's Note: Presented here is a sampling of photographs from Dispersal, a project by writer and photographer Anna Laurent. Dispersal will be on display in the Arnold Arboretum's Hunnewell Visitor Center from October 26, 2013, to January 26, 2014. S eed pods are incredible little vessels tasked with protecting seeds as they mature, and assisting with their dispersal. Dispersal explores how species have evolved different forms to fulfill these common functions. Individually, each photograph is a fine art portrait of a unique botanic specimen; as a series, the collection becomes a visual and scientific inquiry into the diversity of botanic design. Each specimen profile includes text exploring the seed pod's relationship to its parent plant and natural environment, and a bit about its ethnobotanical history. The project explores questions of structure and behavior: Why are the magnolia's seeds red? Why do lotus seeds remain dormant for so long? Why do some trees hold onto their fruits until the seeds have dispersed, while others release the entire vessel? The project began in Southern California, where I began collecting seed pods--dangling from tree canopies and lodged between sidewalk cracks--in my urban Hollywood neighborhood. I have since collected specimens in the rain forests of Hawaii, the deserts of northern Iraq, and ecologies throughout the United States, including the Arnold Arboretum. Seed structures were collected from sites as lush as Lotusland in California (seen here, Nelumbo nucifera) and as sere as the foothills of Kurdistan. All photos by the author. 26 Arnoldia 71\/2 Dispersal 27 Oriental poppy (Papaver orientale) Collected: September 2010 in Portland, Oregon European plant collectors gathered seeds from mountains adjacent to the Euphrates River in the early eighteenth century, and the species has since dispersed in temperate regions throughout the world, including the Pacific Northwest and gardens in Portland, Oregon. With its tomato-red petals and eggplant-colored anthers, the Papaver orientale blossom has long been a darling of botanical illustrators. It is beautiful. The seed pod, however, better reveals the unusual morphology common to species in the Papaveraceae family: there is no style, instead a collection of sessile stigmas radiates atop the ovary, now swelling above a tangle of dehiscent anthers. When mature, seeds are released through small apertures that open below the stigmatic disc. The puckered pod shivers in the breeze, tossing small black bits like a pepper-pot. 28 Arnoldia 71\/2 Dispersal 29 Fleshy-flowered Spindletree (Euonymus carnosus) Collected: October 2012 at the Arnold Arboretum in Boston, Massachusetts In the fall, the Arnold Arboretum's grove of Euonymus trees is a pink haze of plump ornaments. Trees in the genus are characterized by their inconspicuous green or yellow flowers, while the fruits are brightly-colored capsules that split open to reveal seeds covered in an equally bright arillus. Like many other species in the genus, Euonymus carnosus appeals to its primary seed dispersal agent--birds-- with small flashes of red fleshy skin, from which the black seeds seem to coalesce like droplets. Birds, attracted to red wavelengths, pluck the seeds and consume the wattle-like skin. The seeds are carried for the length of the courier's digestive tract, then dispersed. The specific epithet, carnosus, translates to \"fleshy,\" which refers to the thick flower petals and fruits. Euonymus means \"aptly-named,\" an etymology with origins in Greek mythology. The genus of fiery-leaved trees was believed to descend from Euonyme, the mother of the Furies, as the fruits of the tree are beautiful but can be poisonous. Euonymus carnosus is native to forests and woodlands in parts of China and Japan; multiple accessions of this species have grown well at the Arnold Arboretum. 30 Arnoldia 71\/2 Dispersal 31 Astragalus Collected: June 2011 near Jarmo, Kurdistan, northern Iraq It's not always easy--or possible--to identify a seed pod collected in the wild. I found these in the Iraqi steppe near Jarmo, a neolithic farming site in Kurdistan's foothills, one of the first settlements to cultivate grains. I was working on a documentary about the future of agriculture in the Fertile Crescent, and, as we were finishing an interview with an antiquities curator, I set down my camera to collect these capsules. They would prove difficult to identity. One can often refer to a flora (a manual listing all plants in a country), but Iraq lacks this document. Begun years ago, it was delayed due to wars and sanctions. Botanists from Iraq and the Royal Botanic Garden Edinburgh have recently resumed the project, but it will take a while. Having yet to collect this species in Iraq's vast desert wilderness, none of the botanists were able to identify the pods. Meanwhile, I've tentatively named it an Astragalus and have sought confirmation from an Astragalus expert in neighboring Iran. When he replies to my query, I'll have a line to contribute to Iraq's flora. Anna Laurent is a flora-focused writer, producer, and photographer. "},{"has_event_date":0,"type":"arnoldia","title":"Standing Tall: The Upright Swede (Tilia cordata 'Swedish Upright')","article_sequence":5,"start_page":32,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25561","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24e8926.jpg","volume":71,"issue_number":2,"year":2013,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Standing Tall: The Upright Swede (Tilia cordata `Swedish Upright') Michael S. Dosmann S ome fifty years ago, the Arnold Arboretum introduced a cultivar of littleleaf linden (Tilia cordata) to the world. In the same issue where the Arboretum witchhazel introduction Hamamelis "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23437","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170ab6e.jpg","title":"2013-71-2","volume":71,"issue_number":2,"year":2013,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Mutants in our Midst","article_sequence":1,"start_page":2,"end_page":14,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25555","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24eb76d.jpg","volume":71,"issue_number":1,"year":2013,"series":null,"season":null,"authors":"Friedman, William E.","article_content":"William E. Friedman W hat is horticulture? At its core, it is a human celebration, whether conscious or unconscious, of the very fact of evolution. It is thousands of years of detecting and rejoicing in the rare: the selection of the novel form that somehow pleases the human aesthetic or serves to feed the world. Although often overlooked, many of the wonderful horticultural varieties that grow in botanical gardens (as well as in backyard gardens) are premier examples of the amazing and ongoing process of evolution: random mutations that lead, on the rarest of occasions, to novel and desirable biological characteristics--as opposed to novel and neutral or undesirable characteristics. Charles Darwin was an avid consumer of horticultural literature and information, and was a frequent correspondent with the most eminent horticulturists of the nineteenth century. Over the course of his life, he wrote 55 notes and articles in the Gardeners' Chronicle and Agricultural Gazette, one of the most widely circulated horticultural periodicals of his time. He covered everything from how pea and bean flowers are pollinated (Darwin 1857, 1858, 1866) to the origin of variant forms of roses in cultivation (Darwin 1868). He wrote of his observations of and interest in the origin of double-flowered forms (Darwin 1843) and variegated leaves (Darwin 1844). No horticultural phenomenon was beyond his interest. Indeed, Darwin looked to the world of horticulture and plant domestication in order to gain critical insights into the generation of variation and the process of natural selection that underlie evolutionary change. In essence, Darwin was intensely interested in mutants in our midst. Charles Darwin wrote about many horticultural topics including variegated pelargoniums, which were very popular in the Victorian era. `Mrs. Pollack', seen here, was introduced in 1858. \"Florists have attended in some instances to the leaves of their plant, and have thus produced the most elegant and symmetrical patterns of white, red, and green, which, as in the case of the pelargonium, are sometimes strictly inherited.\" -- Charles Darwin, The Variation of Animals -- and Plants Under Domestication, 1868 Evolution at thE arborEtum The Arnold Arboretum of Harvard University hosts a remarkable collection of more than 15,000 accessioned woody temperate plants distributed in over 2,000 different species. This living collection contains wild-collected trees, shrubs, and vines, as well as a spectacular set of horticultural varieties whose very presence is the result of human discovery and propagation of desirable variants. Many of these horticultural varieties are the result of the never-ending process of spontaneous mutations that occur in all organisms and serve to create novel GeOff BryANT mutants in our midst variety, Form, or Cultivar? how to namE a variant plant is the topic of some taxonomic debate and often results in multiple versions of the plant's name. as taxonomic understanding and interpretation changes through the years it often results in changed nomenclature, reflected in the International Code of Nomenclature and the International Code of Nomenclature for Cultivated Plants. a quick reference search finds the white-flowered redbud mentioned in this article listed as Cercis canadensis var. alba, Cercis canadensis f.[forma] alba, or Cercis canadensis `alba'(a cultivar name). the same range of synonyms are found for the mutant Kalmia latifolia (var. polypetala, f. polypetala, or `Polypetala') featured later in the article. to add to the confusion, in common usage the words \"variety\" and \"form\" are often broadly applied (\"i like pink varieties of roses\") or used when referring to a cultivar. For this article, i have used the scientific names as they appear in the arboretum's collections database. Eastern redbud (Cercis canadensis) blooms throughout its canopy, producing a spectacular spring display. traits--the very stuff of evolution. These variant plants, referred to as \"sports,\" arise in a single generation and have undergone a dramatic change in phenotype (the biological properties of the organism) from the parent plant and species. Typically, sports are discovered as a single branching system on a tree or shrub that differs significantly in its morphology, coloration, or other biological properties from the rest of the parent plant. The source of the biological novelty is random mutation, and subsequent vegetative propagation (e.g., grafting, rooting of cuttings, tissue culture) allows the new form to be cloned for further dissemination. Since arriving at the Arnold Arboretum in January 2011, I have fallen in love with these wonderful horticultural results of random genetic mutations and the creation of novelty PAUl W. Meyer, MOrrIS ArBOreTUM 4 Arnoldia 71\/1 Mutants in our Midst 5 WIllIAM e. frIeDMAN WIllIAM e. frIeDMAN This cluster of flowers shows both the normal (pink and magenta) and the mutant (whitish) forms found on the Arboretum's mutant redbud. mostly, but not entirely, white. It was not, however, until the spring of 2009 that these aberrant flowers were first noticed by Arboretum staff. The flowers are beautiful, and novel and rare in a way that every lover of new horticultural forms can appreciate. Now, every year, this redbud continues to produce the typical pink and magenta flowers on most of its shoot systems, with whitish flowers on a single lateral set of branches that bear the mutant gene that results in altered flower color. Interestingly, this is by no means the first horticultural variant of the eastern redbud to sport white flowers. A widely grown one, Cercis canadensis `Alba' (often referred to as C. canadensis f. alba from its earlier botanical description) can be found in gardens throughout the United States. It has pure white flowers, with no trace of red pigmentation. Although it has not been scientifically studied, it is very likely that the mutation that created `Alba' was one that \"broke\" or entirely suppressed the expression of the biochemical pathway to produce red pigmentation in these plants. even young leaves, which typically have a purplish (Top to bottom) normal redbud flower with full red pigmentation; one of the mutant flowers, with pink splotches on the petals and a lighter pink calyx showing that some red pigmentation is still expressed; and a flower of `Alba', with distinctly green sepals and white petals lacking any red pigmentation. 6 Arnoldia 71\/1 Mutants in our Midst 7 of pigmentation that help insect pollinators orient properly as they approach the flower during pollination. Nectar guides are much the same as the lighting on an airport runway, helping the airplane pilot to properly approach the landing strip. finally, in the mutant redbud flowers the female reproductive parts, particularly the style and stigma, differ in pigmentation from the wild type. In normal redbud flowers, the style displays a reddish color, as a consequence of the expression of the biochemical pathways to create anthocyanins. Under the microscope, it becomes evident that the mutant flowers have styles that lack any obvious red pigmentation. What does all of this mean? It suggests that unlike `Alba' and `royal White', which appear to have entirely lost the ability to create anthocyanins (at least in the flowers and young leaves), the Arboretum variant has a mutation that alters where the anthocyanins are produced. In other words, it still makes red pigmentation, but the cellular machinery that might otherwise produce this pigmentation throughout the petals and the style is no longer turned on in these places. How do we know when and where this remarkable single mutation occurred in the Arboretum redbud variant? The answer lies in a basic knowledge of how plants grow and a specific knowledge of an unusual pattern of flowering that can be found in redbud trees. At the tip of every branch of every tree, there is a small group of cells that remains perpetually embryonic and undifferentiated. These cells form the apical meristem, and are similar to stem cells in humans. every year this small population of cells divides, and in dividing creates the new tissues that will differentiate into stems and leaves. If a mutation occurs in one of the cells of the apical meristem, this mutation may come to populate some or all of the cells, and hence the differentiated stem, leaf, and flower cells that are descended from this mutant apical meristem. In the Arboretum's mutant redbud, the mutation that reduced the production of anthocyanins in the flowers of this tree can be found on a set of branches that are all descended from an original mutant meristem of the growing tip of a single shoot. The ability to determine when this mutation occurred in a shoot apical meristem can be deduced because of a specific and somewhat unusual characteristic of all redbud trees. redbuds exhibit a phenomenon known as cauliflory (Owens et al. 1995). Translated literally, cauliflory means flowering on stems. However, in botanical usage, cauliflory refers to the production of flowers on older woody stems. A careful examination of redbud trees reveals WIllIAM e. frIeDMAN This banner petal of a mutant flower clearly shows magenta lines that act as nectar guides for insects (a close-up of the nectar guides under the compound microscope is seen at right). 8 Arnoldia 71\/1 tain laurels produce an abundance of flowers in terminal panicles. In the wild, flowers of Kalmia latifolia are white to pink, with showy cup-shaped corollas. Hundreds of cultivars have been selected; these variants have flowers ranging from white to deep red, many with banded or speckled patterns. But, the \"monstrosity\" described above (initially as Kalmia latifolia var. monstrosa, later as K. latifolia f. polypetala, and now generally referred to as the cultivar `Polypetala') is not a color mutant. rather, it is a variant with an altered morphology of the petals. Instead of forming a sympetalous (fused sets of petals) corolla, `Polypetala' has narrow, unfused individual petals. This is the form of mountain laurel first described by Harvard Professor of Botany Asa Gray in 1870, as a consequence of the keen collecting eye of one Miss Mary Bryant of South Deerfield, Massachusetts. It did not take long before specimens of this unusual morphological mutant came to Harvard University. A specimen of Kalmia latifo- In this inflorescence of Kalmia latifolia `Polypetala' many of the flowers have yet to open. The dark red coloration at the tips of the filiform petals is associated with the unusual production of pollen-producing anthers on these mutant petals. Also note the reflexed normal stamens jutting out between the petals. NANCy rOSe Inflorescences of Kalmia latifolia `Polypetala' create a markedly altered and attractive appearance when the plant is in flower (the plant seen here is the original 1885 accession from South Deerfield, Massachusetts). Flowers of a normal (\"wild-type\") K. latifolia are seen at far left in the photo. Kyle POrT 10 Arnoldia 71\/1 GlASS MODelS: THe WAre COlleCTION Of GlASS MODelS Of PlANTS, HArvArD UNIverSITy; rOSe lINCOlN\/ArNOlD ArBOreTUM Mutants in our Midst 11 lia `Polypetala' from the Harvard University Herbaria notes that it was collected in the Botanic Garden at Harvard (in Cambridge) in 1884. Another 1891 herbarium sheet in the Harvard University Herbaria comes from a grafted specimen that was introduced into the Arnold Arboretum in 1885 (accession 2458). finally, and quite wonderfully, one of the extraordinary models in Harvard's famed glass flowers (formally, the Ware Collection of Glass Models of Plants) was based on observations and collections of the Arboretum specimen of Kalmia latifolia `Polypetala'. In the summer of 1895, rudolph Blaschka--of the father (leopold) and son (rudolph) team that created the glass flowers--came to the Arboretum to sketch and observe this mutant pioneer. The glass model of Kalmia latifolia `Polypetala' (one of over 800 models created by the Blaschkas between 1886 and 1936) can be viewed at the Harvard Museum of Natural History. And, after all of these years, six of the seven original living plants from the 1885 accession (2458-A, B, C, e, f, G) still survive and thrive on the grounds of the Arboretum. In 1907, another cluster of mountain laurels with unfused petals was found along roadsides in leverett, Massachusetts, near Mount Toby (Stone 1909). The mutant petals of these plants were reported not to produce anthers at their termini, as is the case with the `Polypetala' discovered by Miss Bryant and first described by Asa Gray. Arboretum botanist Alfred rehder suggested that this discovery was evidence of the independent origins of these petal mutants in different naturally occurring populations (rehder 1910). However, it is possible that this description was in error. In the University of Massachusetts Herbarium, there are six specimens of the `Polypetala' form of mountain laurel (in flower) that were collected between 1910 and 1932 on Mount Toby, and all of them show anthers at the tips of the mutant petals. Perhaps these oddly placed anthers were not initially observed in the report from 1909. Nevertheless, it is worth noting that `Polypetala'-like forms of Kalmia latifolia have also been found growing in the wild in North Carolina (ebinger 1997) and elsewhere. These variants appear to be fundamentally different from those of the South Deerfield and Mount Toby populations, as they are reported to lack anthers on the tips of the unfused (apopetalous) petals. Clearly there are at least two different and independently formed (evolved) variants with the unifying feature of forming unfused petals--not unlike the multiple evolutionary origins of white-flowered redbuds. Asa Gray's description of the `Polypetala' type of Kalmia refers to the notion that the petals have been \"transformed into stamens.\" In evolutionary terms, this is a statement worth examining. Close observation with a hand lens (or under the microscope) of the \"petals\" of the South Deerfield plant reveals that each one bears a pair of pollen-producing structures at its distalmost end (collectively, an anther). As might be expected, pollen can be found within and then dispersed from these anomalous anthers. Normally, the stamens of Kalmia latifolia comprise a long filament terminated by a reddish anther that produces pollen. A defining characteristic of the floral biology of Kalmia species is that the ten stamens insert themselves into ten pouches in the petals of the cup-like corolla, creating a mechanical tension. visitation by an insect pollinator trips the catapult and the anther flings pollen with enough force to throw it three to six inches away from the flower, but usually directly onto the body of the pollinator, where it will be transported to the next flower to effect pollination (ebinger 1997). In the `Polypetala' Kalmia from South Deerfield, the \"petals\" still produce a pouch about midway along the length of the organ. However, the disruption to the normal morphology of these flowers precludes the proper insertion of the ten normal stamens into these pouches. Thus, as the flower expands towards anthesis (the opening of the flower), the ten normal stamens proceed through their typical pattern of physical reflexing, but never find the petalborne pouches. The \"petals\" also bear much of the typical pinkish-red markings that create some of the brilliant spots or circumferential bands on the corolla of normal flowers. As such, the South Deerfield `Polypetala' \"petals\" may best be thought of as chimeric organs--part petal and part stamen--while some of the other `Polypetala'-like variants that lack anthers on their unfused petals may best be viewed as Mutants in our Midst 13 family history and modern genetics make clear that the gene for hemophilia did not exist in her family prior to her conception. Mutations happen in gametes (or gamete-producing cell lineages), and zygotes and the organisms that develop from the act of fertilization will exhibit the consequences of the new mutation. recent sequencing of whole genomes of human families indicates that each of us carries roughly 75 new simple genetic mutations (\"single nucleotide variants\" in the parlance of geneticists) that neither of our parents was born with (Campbell et al. 2012; Kong et al. 2012). Whether the mutation that created a new chimeric corolla form in the South Deerfield Kalmia latifolia took place in the immediate decades before Miss Bryant found the monstrous plants, we will never know. It could be that this mutation was present in this local population of mountain laurels for hundreds if not thousands of years, unseen by human eyes. And for all we know, this mutation might ultimately mark the beginning of a new species of Kalmia over the course of time. In either case, it took a wandering (and observant) naturalist to discover this product of the evolutionary process, this biological gem, and bring it to the attention of a professional botanist. One can only imagine the delight of Miss Bryant upon finding this unique type of mountain laurel! Botanical gardens are filled with examples of spontaneous mutations, many of which evolved and were discovered in our own lifetimes. These are the very same kinds of mutations that occur constantly in nature and have served as the raw materials that gave rise to humans, oak trees, and plasmodial slime molds--all descended and transformed over the course of billions of years from a single-celled common ancestor of all of life on earth. The raw ingredients of evolution writ large are all around us. And if we look carefully, we can observe the process of evolution by simply walking through a botanical garden, or one's own backyard. Mutant forms of redbud and mountain laurel, as well as myriad other \"sports,\" are an important reminder that we live in a beautiful and profoundly evolutionary world. References Anonymous. 1922. White red-bud. Missouri Botanical Garden Bulletin 10(6): 110. Campbell, C.D. et al. 2012. estimating the human mutation rate using autozygosity in a founder population. Nature Genetics 44: 1277 PostsCriPt: One question that lingered after all of the historical research on Kalmia latifolia `Polypetala' was whether any of the mutant plants (or their descendants) that were originally found on Colonel Bryant's property were still in existence. A map of the South Deerfield, Massachusetts, area from 1871 showed exactly where this property was located. fortunately, this map could be cross-correlated with modern maps to show where Miss Bryant collected the mutant plants. On June 22, 2013, I drove to South Deerfield to hunt the wild mutant Kalmia. The old home that once belonged to Colonel Bryant still stands and is well cared for. regrettably, the land around the original six acres has not had a kind interaction with humans. The barren area on the other side of the brook was home to a pickle factory for many years. The town also installed a major sewer line that is buried alongside the brook. While I found lots of poison ivy and a modest amount of undergrowth beneath some maples and hemlocks, there were no Kalmia plants, mutant or otherwise, to be seen. After my visit to South Deerfield, I drove around the base of Mount Toby. There, I spotted several spectacular populations of mountain laurel in full bloom. My ramble in the woods did not turn up any mutant flowers. Next year, with a bit of time and coordination with the University of Massachusetts Herbarium, we will try to explore the Mount Toby area and search more This section from an 1871 map of South Deerfield, thoroughly for the `Polypetala' form of Kalmia latifolia. Massachusetts, shows Colonel Bryant's property, The loss of the mountain laurel population from where the mutant mountain laurel was discovered, near the center. which Miss Bryant collected the `Polypetala' mutant is a stark reminder of the incredible importance of botanical gardens as refugia for rare and endangered plants, whether entire species, threatened local populations, or unusual mutant forms. It is a very fortunate thing that Miss Bryant's monstrosity was propagated and cared for at the Arnold Arboretum. Otherwise, it might well have disappeared from the face of the earth without a second thought. Kong et al. 2012. rate of de novo mutations and the importance of father's age to disease risk. Nature 488: 471"},{"has_event_date":0,"type":"arnoldia","title":"Rediscovering Rhododendron Dell, Part 2","article_sequence":2,"start_page":15,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25556","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24ebb26.jpg","volume":71,"issue_number":1,"year":2013,"series":null,"season":null,"authors":"Port, Kyle","article_content":"Rediscovering Rhododendron Dell Part 2 Kyle Port \"They [hoodlums] deliberately twist off the metal labels from trees and shrubs, so that valuable information is sometimes lost forever and the yearly replacement bill is terrific. They break hundreds of unopened flower buds off the Rhododendrons in the early spring.\" --Edgar Anderson, Arnold Arboretum arborist , June 4, 1932 All Images By The Author unless otherwise indicated Planted in close proximity to one another, Rhododendron 'Old Port' 990-56-B (a catawbiense hybrid with \"vinous crimson\" flowers, seen here) was incorrectly labeled as R. 'Red Head' 329-91-A (with \"orient red\" flowers). A description published by the Royal Horticultural Society was used to verify the only remaining plant as 'Old Port'; a lack of indumentum on the undersides of the leaves distinguishes it from 'Red Head'. The Arboretum's plant records attest to episodes of vandalism, arson, theft, and other willful shenanigans that have occurred in the living collections over the years. In 2010, a pile of plant record labels was found in Rhododendron Dell. This intentional--and completely unsanctioned--removal of labels from numerous specimens by an anonymous person(s) can certainly be considered a major transgression. But, to quote Albert Einstein, \"In the middle of difficulty lies opportunity,\" and this act of vandalism initiated an unplanned curatorial review that has advanced our understanding of the rhododendron collection and further fostered its use. In response to the identity crises in Rhododendron Dell, a multi-year collection review was conceived. Identity verification and field work (e.g., labeling, photographing) was timed to coincide with peak flowering. Winter months were dedicated to auditing and digesting the raft of secondary documentation (e.g., records, articles, herbarium specimens, images) amassed over the collection's 141-year history. Through 16 Arnoldia 71\/1 Rhododendron Dell, Part 2 17 Labeling Following the imaging and field checks, hundreds of new anodized aluminum records labels were embossed and placed in Rhododendron Dell. many are mounted on three-inch stainless steel screws at the base of large stems. Additional records labels have been hung on branches for easy retrieval. In addition to these, prototypes of larger photo-anodized aluminum display labels were tested over the peak flowering periods. Feedback regarding these labels has been overwhelming positive and the roll-out of permanent signage is expected in 2014. Mapping The current maps of Rhododendron Dell are being revised. Vector data (e.g., points, lines, and polygons) representing plants and hardscape features are being re-collected using global posistioning system (gPs) equipment. These technologies allow for decimeter-accurate field mapping and update the triangulation and submeter-accurate data collection of the past. note that interactive maps of Arboretum collections are available at http:\/\/arboretum.harvard.edu\/plants\/collection-researcher\/ The gorgeous cultivar 'Brookville' was introduced in 1959 by the Westbury Rose Company based in Long Island, New York. Winter Audits and Records Review Nomenclatural review In advance of label production, we undertook a comprehensive review of rhododendron nomenclature. A total of 103 cultivar names were standardized following The International Rhododenron Register and Checklist (Royal horticultural society 2004). This effort revealed inaccuracies in spelling, punctuation, and use of synonymy for 20 elepidote cultivars. In addition to these edits, the name records in BgBase (collections management software) were appended with hybridizer, introducer, parentage, awards, descriptions, and common name as found in the aforementioned resource. We have used this information to create new display labels and have updated online resources. Archival maps and records The first maps documenting the location of accessioned plants in the permanent collections were purportedly authored by henry sargent codman in 1887. Plan views of the landscape On larger specimens, new record labels have been attached to lower trunks with screws. This specimen of R. 'Purpuream Elegans', accession 6135-B, came to the Arboretum in 1891 from the nursery of Anthony Waterer, who hybridized this and many other rhododendron cultivars. Arnold Arboretum from this era were copied from the Frederick law olmsted papers in 1987 but as yet do not reveal individual planting sites. Fortunately, the detailed cartography begun by le Rhododendron Dell, Part 2 19 Rhododendron flower color is often lost in herbarium specimens; compare the 1936 specimen of 'Melton' (left) to a current digital image of its flowers (right). In 2010, grant funds awarded through the museums for America program of the Institute of museum and library services (Imls-mFA) allowed Jonathan Damery, then a curatorial assistant, to scan and georeference the collection of hand-drawn maps. using ArcgIs software, these rasters can be layered with current representations of the Arboretum grounds. In addition, they can easily be printed on 11- by 17-inch paper for problem solving in the field. The Imls-mFA grant also provided resources to enter the Arboretum's entire plant records card catalogue and review accession books (dating from 1872 to 1987). spearheaded by curatorial assistant Kathryn Richardson, the entry of these data has improved all aspects of curatorial work. Herbarium resources A curatorial review would not be complete without a thorough review of specimens in the Arboretum's cultivated herbarium. In the case of hybrid rhododendron, these resources are limited for one major reason: flower color. often lost in the drying process, flower color variations (including the blotch on the dorsal lobe) are critical identification characters of rhododendron hybrids. other flower data such as truss height, width, shape, fragrance, and number of buds can be difficult to discern (or be entirely absent) from a two-dimensional dried specimen. Without question, examination of the whole plant at relevant phenophases provides a more accurate determination. The importance of identifying rhododendron flower color accurately is well documented. Arboretum horticulturist Donald Wyman was a proponent of hte nickerson clor Fan published by the American horticultural society and used this resource to describe the flowers of Rhododendron Dell collections (Wyman 1969). Agents of the Royal horticultural society, United Kingdom, have also published a color chart, which many have used to describe rhododendron cultivars (leslie 2004). These color designations have been saved to the Arboretum's plant records database and are easily retrieved. 20 Arnoldia 71\/1 other notable cultivars as of January, 2013 Subsection Species Traits Valued By Hybridizers Large flowers (some of the largest of the genus) Scented flowers; heat resistant Extreme hardiness; tolerant of exposed sunny sites Tolerant of poor, dry soil Flowers often with crinkled lobes, rachis fairly tall Large, narrow, dark green leaves Species commonly used as understock Hardiness; thick indumentum Hardiness; early flowering Hardiness; early flowering Small stature; longevity of leaf retention Hardiness Tolerant of dry soils Leaf, silvery indumentum; flower bright red to carmine, rarely pink or white Leaf, rusty brown indumentum; flower with purple spotting in throat Hardiness (variable) NATIVITY % of total (RD) cultivars (n = 103) with known parent (backcrosses not tallied) Fortunea Fortunea R. griffithianum R. fortunei E. Nepal, Sikkim, Bhutan, N.E. India Most widely distributed Chinese species. E. United States; Southeastern Appalachian Mountains N.E. Turkey and parts of the Caucasus W. North America 3% (n = 4) 7% (n = 8) Pontica R. catawbiense 48% (n = 50) Pontica R. caucasicum 2% Pontica R. macrophyllum < 1% (n = 1) Pontica Pontica Pontica R. maximum R. ponticum R. smirnowii E. North America Caucasus and N. Turkey N.E. Turkey and Caucasus E. Russia, Siberia, Mongolia, N. China, Japan E. Siberia, China, Mongolia, Korea, Japan China: W. and N.W. Yunnan E. North America China Himalayan foothills, Kashmir to Bhutan 5% (n = 6) 5% (n = 6) 2% (n = 3) Rhodorastra R. dauricum 1% (n = 2) Rhodorastra Neriiflora Pentanthera Scabrifolia R. mucronulatum R. haematodes R. prinophyllum R. racemosum R. arboreum ssp. arboreum R. arboreum ssp. cinnamomeum var. roseum (Album Group) R. ciliatum 2% (n = 3) < 1% (n = 1) < 1% (n = 1) 1% (n = 2) 2% (n = 3, two are R. arboreum) Arborea Arborea E. Nepal, N.E. India, Bhutan, S. hardiest indumented rhododendron species. Its distinctive indumentum and crinkled petal edges are traits favored by hybridizers. References cox, P.A. and K.n.E. cox. 1997. The Encyclopedia of Rhododendron Species. Perth, scotland: glendoick Publishing. leet, J. 1990. The hunnewell Pinetum: A long standing Family Tradition. Arnoldia 50(4): 32"},{"has_event_date":0,"type":"arnoldia","title":"The World of Mosses","article_sequence":3,"start_page":26,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25557","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24ebb6b.jpg","volume":71,"issue_number":1,"year":2013,"series":null,"season":null,"authors":"Stuber, Stephanie","article_content":"The World of Mosses All IMAges By The AuThor Stephanie Stuber W hile the more charismatic trees and flashy flowers initially catch our attention, mosses have an enchanting, charming presence. What is it about these tiny plants that intrigue us? Perhaps we are aware that there is so much more to their story, but their secrets remain intangible, concealed by their diminutive size. Mosses differ from other plants largely in their life cycle. Mosses and tracheophytes (traditionally known as vascular plants) both alternate between two conditions throughout their lives, the gametophyte and sporophyte. The gametophyte is haploid (n), having 1 set of chromosomes, and the sporophyte, being the product of fertilization, is diploid (2n) with 2 sets of chromosomes. Tracheophytes conceal their gametophytes in reproductive structures, like flowers, never to be seen while they develop into the gametes (sperm and egg). For tracheophytes, the dominant condition is the sporophyte--the woody or herbaceous plant itself. The sporophyte produces spores that remain hidden when they develop into the gametophytes which then develop into the gametes. In essence, the gametophytes are dependent on the sporophyte. Mosses carpet the forest floor at the Coastal Maine Botanical Gardens. But in mosses, the sporophyte is dependent on the gametophyte. The domgametophytes produce gametes and the sporoinant condition is reversed; the conspicuous phyte produces spores. The spores, however, are green leafy plant is the gametophyte, and the released into the air before they develop into sporophyte is an ephemeral structure produced the gametophyte, rather than remaining hidden seasonally. The roles are the same, though-- in reproductive structures. Mosses 27 9 10 7 8 1 2 3 6 5 4 Sexual RepRoducTion in MoSSeS 1. a leafy female gametophyte (n) with attached terminal sporophyte (2n). 2. a papery protective covering, the calyptra (n), sheds off when the capsule (2n) fully develops; a remnant of the interior archegonial wall. Spores develop by meiosis inside the capsule. a row of tiny teeth, the peristome (2n), aids in spore dispersal. a spore (n) settles on a place to germinate. The protonema (n) emerges from the spore, reminiscent of filamentous algae, and develops into mature male and\/or female plants depending on the species. 3. 4. 5. 6. a cap, the operculum (2n), pops off the capsule when spores are mature. 7. a cluster of antheridia (n) develop on the male. 8. a cluster of archegonia (n) develop on the female. 9. in the presence of water, flagellate sperm (n) are released from the antheridium and swim to the egg (n) in the archegonium to fertilize it. 10. The fertilized egg (2n, zygote) develops inside the archegonium and emerges as the sporophyte. 28 Arnoldia 71\/1 Mosses 29 higher plants today. This model may also bridge the gap between their aquatic algal ancestors and the terrestrial tracheophytes. MoSSeS up cloSe When you first take a look at a moss plant, with your naked eye or under a hand lens, often the first thing you notice are striking similarities to other plants. Mosses have stems, tiny leaves, and little rootlike structures. With the aid of a microscope you may see more parallels: a midrib, a serrated margin, conductive tissues, even tomentum. These structural analogs have similar purposes in both mosses and tracheophytes. Mosses come in an enormous array of shapes, sizes, forms, colors, and textures, but most are made up of the same components. Members of the genus Polytrichum are commonly used to represent a typical moss species because of their relatively large size and distinct features. The gametophyte consists of parts similar to most other tracheophytes. The leaves of mosses are called phyllids to distinguish them from the true leaves of tracheophytes, which have lignified vascular tissues, but bryologists will call them leaves regardless, understanding their technical differences. These simple leaves are arranged spirally along the stem. This is a good distinguishing characteristic from liverworts, whose leaves are distichous (arranged in a two-ranked fashion on opposite sides of the stem). Instead of roots, mosses have similar structures called rhizoids. They do not make up an extensive subterranean network; rather, they are superficial and act more as a holdfast to anchor the moss to its substrate. The sporophyte consists of a stalk called a seta and the capsule, whose main parts are shown in the lifecycle image. My undergraduate professor, Dr. robin Kimmerer, described mosses as \"time made visible,\" and mosses undoubtedly do lend a certain timeless aesthetic to the landscape. Intuitively we relate the amount of mosses in an area to the length of time it has remained undisturbed. What perpetuates their reputation for being slow growing? Mosses, unlike most life forms on this planet, are poikilohydric. This means that they cannot internally regulate water, so are subject to moisture fluctuations in their The acrocarpous Ulota hutchinsiae has sporophytes that emerge terminally from the gametophyte. The mat-forming pleurocarp Hypnum imponens sends out sporophytes laterally. 30 Arnoldia 71\/1 Mosses 31 filamentous and essentially formless, offering much of their surface area to the open environment. As you move through mesic to drier habitats, the forms become more complex. The dendroid forms are still loose, but have rigid stems to support upright growth on saturated land. The pinnate forms with more intricate and rigid designs increase the amount of capillary spaces, helping to conserve water in mesic areas. habitats with limited water tend to support turf and cushion forms best. Their tight, dense forms and specialized cellular structures and appendages facilitate water retention in drier environments. Their desiccation tolerance is also directly related to their morphology; those species that live in wet areas will have less tolerance to desiccation than those species that are subjected to intermittent water availability. Because of their poikilohydric nature, mosses have had to develop ways to survive those dry periods in order to continue colonizing land further away from a water source. The length of time that some mosses can survive without water is remarkable. Aquatic mosses can remain desiccated for a few months to as much as a year, mesophytic species can wait several years without water, and xerophytic species are known to survive decades or centuries without water. once water returns, they will begin repairing the cellular damage incurred by the desiccation process and then begin photosynthesizing once more. of course, this is observed along a spectrum. The trend between form and desiccation tolerance, though positive, is dependent on the rate of the desiccation process; the slower the drying rate, the longer it can survive in that state. A close-up view of sphagnum moss reveals its rich texture. 32 Arnoldia 71\/1 Mosses 33 Polytrichum piliferum gives off a silvery cast with its very long clear awns and thin waxy cuticle that covers the leaves. awns are often white or greyish, which is thought to aid in light reflection, thus cooling the plant and protecting it from damaging ultraviolet light. These awns extend beyond the leaf margins, increasing the boundary layer blocking desiccating air flow. some species have found ways to thicken their leaves to help retain water longer. some can have short protrusions on the cell surface called papillae. Papillose species have a dull, matte appearance from a distance because of their roughly shaped cells, as opposed to the shiny appearance of species with smooth cells. Members of the Polytrichaceae have lamellae--multistratose plates of cells aligned perpendicularly over the leaf surface, effectively thickening the The awn of Tortula ruralis. 34 Arnoldia 71\/1 Mosses 35 A moss microcosm composed of star-shaped Polytrichum commune, windswept Dicranum scoparium, and short, pale Leucobryum glaucum. Crum, h. A. and l. e. Anderson. 1981. Mosses of Eastern North America. 2 vols. New york: Columbia university Press. glime, J. 2007. Bryophyte Ecology. 5 vols. ebook sponsored by Michigan Technological university and the International Association of Bryologists. Accessed in 2013 at http:\/\/www.bryoecol.mtu.edu goffinet, B. and A. J. shaw. 2009. Bryophyte Biology. 2nd edition. Cambridge, united Kingdom: Cambridge university Press. Kimmerer, r. W. 2003. Gathering Moss: A Natural and Cultural History of Mosses. Corvallis, oregon: oregon state university Press. Malcolm, W. and N. Malcolm. 2006. Mosses and Other Bryophytes: An Illustrated Glossary. Nelson, New Zealand: Micro-optics Press. stephanie stuber is a former Arnold Arboretum Curatorial Fellow and author of The Secret Lives of Mosses: A Comprehensive Guide for Gardens."},{"has_event_date":0,"type":"arnoldia","title":"Chamaecyparis obtusa 'Chabo-hiba' 877-37: A Venerable Survivor","article_sequence":4,"start_page":36,"end_page":37,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25554","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24eb728.jpg","volume":71,"issue_number":1,"year":2013,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Chamaecyparis obtusa `Chabo-hiba' 877-37: A Venerable Survivor Peter Del Tredici W hen people ask \"What's the oldest tree growing at the Arnold Arboretum?\" they're usually surprised to learn that it's a 276-year-old compact hinoki cypress (Chamaecyparis obtusa `Chabo-hiba', accession 877-37) that stands only four feet tall. It is one of seven `Chabo-hiba' specimens in the Larz Anderson Bonsai Collection that were imported from Yokohama, Japan in 1913. This makes 2013 a milestone for the tree--the hundredth anniversary of its arrival (and survival) in North America. It makes my head spin to think that someone has been watering this plant pretty much every day since well before the American Revolution! While this `Chabo-hiba' is not the oldest Japanese bonsai in the United States (there are older ones at the United States National Arboretum in Washington, D.C.) the Arboretum's plant has been under continuous cultivation longer than any other bonsai growing in North America. Larz Anderson attended Harvard College (class of 1888) and later served as a diplomat in the Foreign Service. In 1912, near the end of the Taft administration, he was appointed \"Ambassador extraordinary\" to Japan, a post he held for only six months, until Woodrow Wilson moved into the White House. During his brief stay, Anderson was smitten by the \"bonsai bug,\" and in early 1913, shortly before completing his posting, he purchased at least forty plants from the Yokohama Nursery Company to bring back to his estate in Brookline, Massachusetts. Many of the specimens offered for sale by the nursery were already hundreds of years old. Photographs from the time show that the `Chabo-Hiba' plants were often trained into a conical shape-- suggestive of a distant mountain--with regularly arranged, horizontal branches. Anderson and his wife Isabel (Weld) left Japan on March 6, 1913, and it seems likely that the plants followed them across the ocean in a shipment that autumn. Once they arrived, the trees were displayed on the terraces of the Anderson home where they resided for nearly twenty-five years. The collection was donated to the Arboretum in two batches, initially in 1937 following Larz's death, and later in 1948, following Isabel's death. `Chabo-hiba' 877-37 came to the Arboretum in the first installment and was put on display along with the other plants in a lathhouse on the grounds of the former Bussey Institution. They remained there until 1962 when they moved into their current hexagonal home near the Dana Greenhouses. In 1969 the Arboretum appointed Connie Derderian to take care of the plants. As honorary curator, Connie revitalized the collection after years of neglect and took care of the plants until 1984. Having worked as Connie's apprentice since 1979, I became the new curator the year she retired. In 1998, the noted English bonsai master, Colin Lewis, became involved with the collection. The fact that seven large `Chabo-hibas' have survived the ravages of both time and occasional neglect for the past hundred years is a testament to the incredible durability of the plants themselves. By virtue of their longevity, the plants provide a direct link not only to the early 1900s, when wealthy Americans were passionately collecting cultural artifacts from Asia, but also to the Tokugawa era in Japan (1600 to 1868) when shoguns ruled the land and the plants themselves occupied places of honor in temples throughout the country. The hinoki cypress cultivar name chabo-hiba is not widely used in Japan today, and it took some effort to uncover its history and meaning. The word hiba is the common name for the arborvitae-like conifer Thujopsis dolobrata and means \"hatchet-shaped,\" in reference to the scale-like foliage of the plant. Chabo means bantam or dwarf chicken, and when combined with hiba means \"compact or dwarf cypress.\" In the landscape, Chamaecyparis obtusa `Chabo-hiba' is a relatively slow-growing plant that develops a pyramidal shape when left unpruned. When grown in a container and intensively pruned, it produces congested, planar foliage and contorted horizontal branches, resulting in striking bonsai specimens like accession 877-37. Peter Del Tredici is a Senior Research Scientist at the Arnold Arboretum. Selected specimens from the Larz Anderson Bonsai Collection will be on display at the Isabella Stewart Gardner Museum in Boston from October 2nd to 13th, 2013."},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23436","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170ab28.jpg","title":"2013-71-1","volume":71,"issue_number":1,"year":2013,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Beyond Pine Cones: An Introduction to Gymnosperms","article_sequence":1,"start_page":2,"end_page":14,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25549","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24eab28.jpg","volume":70,"issue_number":4,"year":2013,"series":null,"season":null,"authors":"Conway, Stephanie","article_content":"Beyond pine Cones: An Introduction to Gymnosperms Stephanie Conway G ymnosperms are an intriguing group of plants, yet in many ways they are not well known. Most people can recognize a pine, with its familiar woody cones, but they may not know that this and other conifers are gymnosperms. Or, they may think that conifers are the only plants in the gymnosperm group. Undoubtedly the often large-flowered angiosperms (flowering plants) are the better known group within the seed plants, but gymnosperms are well worth a look. So what are gymnosperms and what makes them so intriguing? There are four groups of plants that make up the gymnosperms: the wellknown conifers, plus the lesser known cycads, ginkgo, and the order Gnetales. These groups KEITH KAnOTI, MAInE FOrEST SErvIcE, BUGWOOd.OrG are so different from each other that it would be hard to immediately recognize them as related. In fact, exactly how they are related to each other is not entirely clear, but most studies put cycads and ginkgo at the base of a gymnosperm evolutionary tree (meaning that they are the simplest, evolutionarily), and conifers and Gnetales as more evolutionarily advanced. What does it mean to be a gymnosperm? The most common feature across all four groups is that the ovule (which becomes the seed) is naked (unprotected) prior to fertilization. In comparison, the angiosperms have ovules that are protected by a layer of tissue called a carpel. The word gymnosperm comes from ancient Greek and means \"naked seed.\" This naked state of the ovule is a unifying feature of the gymnosperms (there are also some shared vegetative features such as wood anatomy), but often these ovules are not visible to the naked eye. This is perhaps what makes them so intriguing: How does this translate to the more common feature that we can see, the cone? How did these evolve? And how does the cone tell the story of the evolution of the gymnosperms? Gymnosperm roots The ancestors of gymnosperms most likely evolved from a group of plants called the seed ferns (pteridosperms), which are known only from the fossil record. These were the first plants to reproduce by seeds, despite looking deceptively like ferns. (True ferns reproduce from spores rather than seeds.) Early seed plants bore their seeds directly on leaves or branches, without any specialized structures like cones. From this starting point we can begin to see how the naked ovules and cones of living gymnosperms evolved. The four lineages of gymnosperms each have a unique set of cone characteristics, and comparisons with the naked eye are extremely difficult. In fact, even comparisons between well-known conifer groups are challenging. To understand Pine cones are perhaps the most familiar gymnosperm cone type. A mature eastern white pine (Pinus strobus) cone is seen here. Gymnosperms 3 STEpAHAnIE cOnWAy Angiosperms Gnetales seed plants Conifers Gymnosperms Vascular plants Ginkgo Cycads Ferns and fern allies Lycopods Phylogeny chart showing the relationship of gymnosperms to other plant groups. the elusive relationship between these cone types, it helps to examine the distinct paths of evolution that each gymnosperm lineage took from the seed fern ancestral condition, how all retained the character of a naked ovule and yet ended up with very different looking reproductive structures. and make them unique among all seed plants. They have a single, typically unbranched trunk with the leaves all bunched together in a crown at the top of the plant. This features makes them look superficially like palm trees, a fact reflected in the common name of one cycad that is often grown as a house plant, sago palm MIcHAEl cAlOnjE, MOnTGOMEry BOTAnIcAl cEnTEr CyCAds cycads are a very ancient lineage of plants with a fossil record that extends back at least 280 million years. They were once very common across most of the planet and were a prominent plant group in the age of the dinosaurs, but they have since retreated to the tropics and sub-tropics. As is the case for all the gymnosperm lineages, it's important to remember that when we look at the cycad taxa growing today we are seeing the survivors of a once very successful plant group. These \"leftovers\" include 3 families of cycads: cycadaceae, Zamiaceae, and Stangeriaceae, which contain about 11 genera and 250 species in total. cycads have unique characteristics that set them apart from the rest of the gymnosperms The female cone of Cycas revoluta. Note that the sporophylls resemble leaves and are all bunched together at the crown, similar to the leaves. Young ovules are formed on the lower portion of the sporophylls and are very exposed or naked. 4 Arnoldia 70\/4 Gymnosperms 5 pETEr dEl TrEdIcI and, as we'll see, shows that the cycad cone and the conifer cone had quite different evolutionary beginnings. But first, let's look at the fascinating Ginkgo biloba, which, in terms of cone morphology, is often considered an intermediate between cycads and conifers. GInkGo Ginkgo biloba is the sole living species of the once widely distributed order Ginkgoales and is often called a \"living fossil.\" This plant has fascinated botanist for centuries because it represents a unique set of characteristics that alludes to both the cycads and conifers but which represents a unique lineage within the gymnosperms. Ginkgo's flat, fan-shaped leaves are its most distinctive feature; the leaves on the plant's long shoots are typically two-lobed, hence the specific epithet biloba. Unlike the cycads, adult trees are heavily branched and have a broad crown. The fertile structures in ginkgo are unique as well, with little to make a comparison to either the cones of cycads or conifers easy. The male pETEr dEl TrEdIcI A Ginkgo biloba tree in fall color at Forest Hills Cemetary in Boston. KEvIn nIxOn The female cones of Ginkgo biloba are generally thought to have evolved from a branch, but all that remain are the long stalks with terminal ovules (seeds) with a thin fleshy covering. Male ginkgo cones (strobili) bear many pollen-producing organs along a central stalk. 6 Arnoldia 70\/4 Gymnosperms 7 umns\" are attached. Each of these attached columns has its own set of organs attached to it. In other words, you can break up a cone into a number of individual units, and each unit has a complete, replicate set of organs. Each one of those units is made up of a bract, a scale, and ovules. The bract is on the outside, and the scale is on the inside. This scale is sometimes called the ovuliferous scale because it is where the ovules are formed and where eventually the seed develops. The fact that the scale where the ovules are formed sits at the base of the bract is important because therein lies the fundamental compound nature of the cone. pETEr dEl TrEdIcI Florin proposed that in the ancestor of the conifers, seeds were formed on widely spaced branches, each branch with a number of fertile scales that bore stalked ovules. Each branch formed at the base of a bract. He proposed that over evolutionary time these branches transformed to have fewer and fewer scales until there was only one, that the ovules lost their stalks, and that the single remaing scale became more and more fused to the bract. So the interpretation is that each unit (an individual bractscale complex) that we break off a cone is all that remains of a once large branch. Most of the other genera in the pine family (pinaceae) have fundamentally the same bractscale complex but with different shapes and sizes of the bracts and scale. In Pinus for example, the bracts are small and inconspicuous compared to the scales, whereas in douglaspETEr dEl TrEdIcI Young female Douglas-fir (Pseudotsuga menziesii) cones sit upright on the branch and display prominent pink bracts (at this stage the scale cannot be seen). The more mature male pollen cones (hanging downward) have pollen organs attached directly to the cone axis. pETEr dEl TrEdIcI Young cone of northern Japanese hemlock (Tsuga diversifolia) with large green and purple scales. The much smaller bracts (white with brown tips) can be seen on the scales closest to the stem. The most prominent feature of this young Sciadopitys verticillata cone is the large white scales, with the smaller brown bracts hidden underneath. 8 Arnoldia 70\/4 Gymnosperms 9 pETEr dEl TrEdIcI The fleshy olive-shaped female cones of Cephalotaxus fortunei. rOBErT vIdEKI, dOrOnIcUM KFT., BUGWOOd.OrG Cones of Taxus (T. baccata is seen here) are so different that they are hard to compare to other conifers. In this species, the seeds are formed terminally on the end of short stems, and a swelling at the base of the ovule develops into a fleshy red aril that covers the seed and also attracts seed dispersers. On the younger green cone the single terminal seed can be seen with the fleshy aril just starting to develop. The large Southern Hemisphere family podocarpaceae also developed a berrylike cone, with fleshy parts to aid dispersal and minimal numbers of seeds per cone. However, this family has a unique cone type that looks nothing like the cones of Juniperus. The cones typically consist of a number of sterile bracts and one fertile bract on which the ovule arises on a structure called the epimatium, which is considered the evolutionary equivalent to the ovuliferous scale. In Podocarpus, the bracts at the base of the cone also swell into an often colorful \"receptacle\" that, as in Juniperus, probably serves in attracting animals for dispersal. plum yew (Cephalotaxus) also has fleshy, single-seeded cones that look suspiciously like olives. The early development of Cephalotaxus shows a lack of ovuliferous scales, and instead the ovules form on the bracts in a manner similar to other conifers. However, the bracts grow out to cover the seed in a fleshy covering that, as seen in Podocarpus, presumably aids in animal dispersal of the seed. Taxus is the final example of a female conifer cone and it's one that does not fit within Florin's theory of conifer cone evolution. The female reproductive structure of Taxus does not have ovules on bracts or scales; instead, it has a single terminal ovule. This ovule sits at the end of a short branch, and an outgrowth at the base of the seed becomes a fleshy red aril that partly covers the seed. Florin himself was so convinced of the fundamentally different nature of the cone structure in Taxaceae that he placed the family in a different order, the Taxales. This implied that Taxales had different ancestors than the rest of the conifers, therefore making the conifers not a natural group. This was a controversial theory, and other researchers have since shown it to be unlikely. Instead, researchers have proposed that the terminal cone may be related to the more advanced cones of the cupressaceae, including various species of Juniperus with single terminal ovules. However, how and from where the Taxus type of cone evolved (if considering the conifers as a monophyletic group) has not yet been satisfactorily resolved and remains something of a mystery. 10 Arnoldia 70\/4 USdA-BlM, USdA-nrcS plAnTS dATABASE Gymnosperms 11 KEvIn nIxOn A male cone of Gnetum gnemon with rings of pollen organs below rings of sterile female ovules, some with pollination drops present. dEnnIS STEvEnSOn, nEW yOrK BOTAnIcAl GArdEn The seed cones on this female Gnetum urens have matured and only one red, fleshy seed has developed from each cone. Above the seed on the right you can see the nodes where the other ovules would have formed, but have failed to develop. Ephedra are generally scalelike, or occasionally longer and needlelike, and all are joined at the base to form a sheath around the stem. Most species of Ephedra are dioecious (separate male and female plants). The pollen cones of Ephedra have a pair of bracts at the base of the cones, and the cones themselves are made up of a series of bracts, each with its own fertile shoot. This makes these cones compound structures in the same fashion as the seed cones of conifers. The female cones are also compound. The cones have a pair of bracts at their base, and the cones themselves are also made up of a series of bracts. The uppermost bracts have ovules in their axes, although often only one develops into a seed. Gnetaceae has only one genus, Gnetum. Most Gnetum species are tropical vines, though one of the most widely studied species, Gnetum gnemon, is a tree. Gnetum species occur in parts of Asia, South America, and Africa as well as some pacific Islands. If you were to walk past one in the tropics you would be hard pressed to recognize it as a gymnosperm because the leaves are broad, flat, and have netlike veins, making it look much more like a flowering plant (angiosperm). Gnetum cones are also very distinct from typical conifer cones and they form fleshy seeds that look like berries. Both the cones that produce pollen and those that produce seeds are compound structures and unique among gymnosperms. In Gnetum gnemon they are long and have distinct nodes where the fertile structures are formed. The pollen cones have bracts that cover the nodes, and underneath these a number of pollen organs are enclosed within two fused structures. Above this ring of pollen organs there are often aborted female ovules, which has lead many botanists to consider the cone of Gnetum to be primitively flowerlike. The seed cone also is on a long axis, with the fertile structures occurring on the nodes. There are bracts that cover a ring of 8 to 10 ovules. Each ovule is surrounded by 3 bractlike structures that form envelopes around the ovule. Welwitschiaceae consists of only one species, Welwitschia mirabilis, which may be one of the strangest plants on the planet. It grows only in the namib desert of Angola and namibia and produces just two huge leaves from a short, woody, unbranched stem. The leaves grow an average of 8 to 15 centimeters (3 to 6 inches) per year, and often are split and twisted at their ends, forming a tangled mass. Some Welwitschia leaves have been measured at up to 6 meters (19.7 feet) long. The plants survive in the desert by developing a huge taproot that may extend down nearly 2 meters (6.6 feet). A few plants have been estimated to be close to 12 Arnoldia 70\/4 Gymnosperms 13 An adult Welwitschia mirabilis plant growing in the Messum River area in Namibia. but the common theme across all the lineages has been an evolution towards simplifying the reproductive structure. This has been achieved in a variety of ways and with different results. cycads reduced the leafy portion of their cones down to a scale. Ginkgo reduced a large branch to a single stalk with two ovules. conifers tended towards simplifying the branch complex to just a bract, or getting rid of the traditional cone altogether, and 4 out of the 7 conifer families developed a fruitlike structure as well as reducing the seed number. Gnetales began experimenting with having both seed and pollen structures within a single cone. While a pine cone may be the best known representative of gymnosperm reproductive structures, it is in fact only a small part of the gymnosperm story. The current, living assemblages of gymnosperm groups are really only relicts of what once was a gymnosperm dominated world, so the task for us is to understand the whole narrative of dominance and decline. The gymnosperms of today are incredibly important since they represent 4 out of the 5 extant lineages of seed plants (angiosperms are the fifth lineage) and botanists continue to study exactly what gymnosperms are and how they evolved. current research includes phylogenetic stud- Male cones of Pinus muricata are simple, with a bract at the base of each cone and the pollen organs attached directly to the cone axis. pETEr dEl TrEdIcI GIllIAn cOOpEr-drIvEr 14 Arnoldia 70\/4 pETEr dEl TrEdIcI "},{"has_event_date":0,"type":"arnoldia","title":"Rediscovering Rhododendron Dell, Part 1","article_sequence":2,"start_page":15,"end_page":18,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25552","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24eaf6b.jpg","volume":70,"issue_number":4,"year":2013,"series":null,"season":null,"authors":"Port, Kyle","article_content":"Rediscovering Rhododendron Dell, Part 1 Kyle Port A pile of plant record labels, mysteriously stripped from accessioned plants in the Arboretum's Rhododendron Dell, sent Manager of Plant Records Kyle Port on a mission to assess, verify, and relabel the collection In this issue, Kyle reports on the early history of Rhododendron Dell, and in the next issue he will write about the multi-layered curatorial process involved in the Rhododendron Dell project. Editorial NotEs rhododendron show at Boston the rhododendron show on the Boston Common was a sight never to be forgotten--the finest in colors and display of choice varieties this country has ever beheld. it was with considerable difficulty the bare privilege was secured from the common council, to exhibit upon the Common; and next, it was believed to be almost impossible to transport and successfully flower so many plants as would be needed to produce the desired effect. thanks, however, to the untiring energy of Mr. H. H. Hunnewell, Charles s. sargent and E. s. rand, jr., every difficulty was surmounted, and for the entire month of June the denizens of that city saw a collection of azaleas and rhododendrons of rare value and great beauty ... two large tents were erected, one about 60 by 100 feet, the other 100 by 300 feet, and the plants transplanted from their native home and conservatories of Mr. Hunnewell and Mr. sargent, and placed in the natural soil of the Common. Within the tents were laid out, first, an avenue of 100 feet in length, bordered with Palms and rare Ferns; this led to the rhododendron beds and walks. in the center of the large tent were three raised beds; the first, 15 by 30 feet; the second, 50 by 80 feet; the third, 15 feet in diameter. Walks also surrounded all the beds, which were lined with specimen plants. imagine all this space and beds filled solidly with masses of rhododendrons in full bloom, bearing flowers of most royal size, and delicate as well as glowing and brilliant colors, and it would seem to be but a vision of the garden of Paradise. ... most of them [rhododendrons] are fit for in-door greenhouse culture only, many being but just imported from the Knapp [Knap] Hill nursery of anthony Waterer, Woking England. luther tucker The Horticulturist, and Journal of Rural Art and Rural Taste, Volume 28, august 1873 O ne hundred forty years ago, a triumphant rho d o d endron show bloomed on the Boston Common. For a nominal fee, attendees were ushered under tents where plants from private collections, including those of Arnold Arboretum director Charles S. Sargent and the event's sponsor, H. H. Hunnewell, were arranged. Rhododendron hybrids imported from Anthony Waterer (Knap Hill Nursery, Woking, England) garnered considerable attention. The revelation that R. `Album Elegans' and a few other cultivars were hardy outdoors in the Boston area soon fostered planting trials beyond conservatory walls, specifically in the bur- 16 Arnoldia 70\/4 ARCHivES OF THE ARNOld ARBORETuM allowed for cold air to sink away toward low-lying Bussey Brook Meadow. The new hybrids were not immediately planted in Rhododendron dell; instead, the first plantings on the site were of the hybrids' North American parent species, which included R. catawbiense, R. maximum, and R. minus. it is likely that the only remaining plants from these early plantings exist in a mass planting of R. maximum accessions 23020 and 23021. These accessions actually comprise a number of accessions that were interplanted over the years and became indistinguishable from each other. The oldest of these R. maximum accessions was obtained as seeds in 1880 from Benjamin Marston Watson's Old Colony Nurseries and Seed Warehouse in Plymouth, Massachusetts. in 1886, the first R. catawbiense hybrids from Anthony Waterer were planted in Rhododendron dell. Some of these hybrid cultivars had been featured in the tents of the 1873 rhododendron show on Boston Common, but now they were being planted outdoors to see how they would fare. Among these inaugural cultivars, R. `Purpureum Grandiflorum' (accession 2804) and R. `Album Grandiflorum' (accession 2805A) survive to this day. Subsequent introductions such as R. `Mrs. Harry ingersoll' (accession 6202-C, acquired in 1891) epitomize the allure these hybrid rhododendrons had upon so many. Their survival at the Arboretum solidified a resolve to develop, evaluate, and maintain a collection for the ages. As Sargent wrote to Anthony Waterer in February 1911, \"i think that we should have here a correctly named standard set of the hardy hybrid Rhododendrons as so many people depend on the Arboretum for information on such a subject.\" While the majority of the early acquisitions of hybrids in Rhododendron dell were those of Anthony Waterer and his cousin, John Waterer, a number of other international hybridizers are also represented. Fel- Rhododendron catawbiense `Grandiflorum' KylE PORT Rhododendron `Mrs. Harry ingersoll' KylE PORT Rhododendron `Echse i' KylE PORT 18 Arnoldia 70\/4 KylE PORT "},{"has_event_date":0,"type":"arnoldia","title":"Book Excerpt: Ginkgo: The Tree That Time Forgot","article_sequence":3,"start_page":19,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25550","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24eab6d.jpg","volume":70,"issue_number":4,"year":2013,"series":null,"season":null,"authors":"Crane, Peter","article_content":"Book exCerpt Ginkgo: The Tree That Time Forgot Peter Crane editor's note: In his new book, noted botanist peter Crane has gathered a vast trove of information on the ginkgo, undoubtedly one of the most loved trees in the world. exploring topics ranging from paleobotany to evolutionary biology, plant exploration, and human culture, the author presents fascinating tales from the ginkgo's very long history on earth. printed here by permission of the publisher is Chapter 21, \"extinction.\" (don't worry, ginkgophiles--Chapter 22 is \"endurance.\") Ginkgo: The Tree That Time Forgot Peter Crane yale University Press, 2013. 408 pages. ISBN: 978-0-300-18751-9 20 Arnoldia 70\/4 Book Excerpt 21 22 Arnoldia 70\/4 Book Excerpt 23 24 Arnoldia 70\/4 Book Excerpt 25 26 Arnoldia 70\/4 Book Excerpt 27 "},{"has_event_date":0,"type":"arnoldia","title":"Ginkgopalooza","article_sequence":4,"start_page":28,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25551","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24eaf26.jpg","volume":70,"issue_number":4,"year":2013,"series":null,"season":null,"authors":null,"article_content":"P GinkGopalooza eter Crane's book inspired us to gather a few ginkgo images. These pages show just a few of the aspects that make ginkgo such a fascinating plant. An ArTfully espAliered ginkgo graces a wall of the sterling Morton library at the Morton Arboretum (right). At about 55 years old, this ginkgo is a youngster compared to the \"old lion\" ginkgo at the royal Botanic gardens, kew. That venerable tree was planted in 1762 and is seen here in an engraving that originally appeared in the British publication Gardeners' Chronicle in March 1889 (lower left), and in a photograph from May 2010 (lower right). Credit photos, clockwise from upper right: Kris Bachtell, Tony Kirkham, Archives of the Arnold Arboretum people love ginkgos. Clockwise from upper left: This wonderful 1921 e. H. Wilson photograph shows a woman hugging an enormous ginkgo at the Temple of the yellow dragon, kuling (su-shan), China (Wilson recorded the tree at 100 feet tall and with a trunk circumference of 19.5 feet, measured at 3 feet above ground level). ginkgos stand behind a Buddha statue at the sens-ji temple in Tokyo. over 100 (possibly 200) ginkgo cultivars have been selected, including those with dwarf, fastigiate, weeping, and globeshaped habits as well as different leaf forms (`variegata' is seen here). ginkgophiles enjoy the golden glow under a ginkgo alle THe ginkgo leAf's simple but elegant shape and unique dichotomous venation make it a work of art in itself. over many centuries its iconic form has appeared on everything from street signs to silverware to shampoo bottles. shown here (clockwise from upper left) are a live leaf, a plate from Japan, the 1916 woodblock print Crow and Ginkgo Leaves by Japanese artist Watanabe seitei, a pair of silver sugar tongs, and a decorative wall tile. Credit photos, clockwise from upper left: Peter Del Tredici, Peter Del Tredici, Smithsonian Institution, Nancy Rose, Kris Bachtell "},{"has_event_date":0,"type":"arnoldia","title":"2012 Weather Summary","article_sequence":5,"start_page":31,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25548","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24ea76f.jpg","volume":70,"issue_number":4,"year":2013,"series":null,"season":null,"authors":"Pfeiffer, Sue A.","article_content":"2012 Weather Summary Sue A. Pfeiffer T emperatures were again above average in 2012; this was the first year in local recorded weather history that all 12 months had above average temperatures (in 2011, all months except for November were above average in temperature). JANUARY began with warm temperatures, but a cold front moved in on the 4th and 5th, dropping the low to 10 32 Arnoldia 70\/4 2012 Weather 33 JULY was the warmest month of the year with highs mostly in the 80s and 90s. The month started off sunny and warm with minimal precipitation and consistently high temperatures in the 80s. a 6-day heat wave (temperatures 90 34 Arnoldia 70\/4 JIM PaPaRgIRIS Arnold Arboretum Weather Station Data "},{"has_event_date":0,"type":"arnoldia","title":"The Sweet Smell of Spring: Abelia mosanensis","article_sequence":6,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25553","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24eb36f.jpg","volume":70,"issue_number":4,"year":2013,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"The Sweet Smell of Spring: Abelia mosanensis Nancy Rose S omeday I really want to publish a scratchand-sniff issue of Arnoldia, providing a \"by the nose\" tour of the Arnold Arboretum. I'd probably skip the less pleasant odors-- skunk cabbage, ripe ginkgo cones, the stinkhorn fungi that pop up in mulched planting beds--in favor of the many truly wonderful scents to be found here. While pleasant fragrances occur from winter (e.g., the flowers of Hamamelis mollis `Princeton Gold') through autumn (e.g., falling Cercidiphyllum japonicum leaves), the floral explosion from mid-spring to early summer brings the peak sniffing season. Come May, there are plenty of sweet-smelling flowers to stick my nose into but my current favorite is the aptly named fragrant abelia (Abelia mosanensis). This deciduous abelia is less well known than glossy abelia (Abelia "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23435","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd170a76e.jpg","title":"2013-70-4","volume":70,"issue_number":4,"year":2013,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Middlesex Fells, a Flourishing Urban Forest","article_sequence":1,"start_page":2,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25546","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24ea326.jpg","volume":70,"issue_number":3,"year":2013,"series":null,"season":null,"authors":"Kittredge, Walter","article_content":"The Middlesex Fells, a Flourishing Urban Forest Walter Kittredge ... five miles northerly of Boston lies a great tract of country, all stony hills and tablelands, almost uninhabited, and of wonderful picturesqueness, and wild, rugged beauty ... The nature of this region cannot better be characterized than by the application of the old Saxon designation fells,--a common enough word in England, meaning a tract of wild stone hills ... Sylvester Baxter, Boston Herald Supplement, December 6, 1879 Bitternut hickory (Carya cordiformis) is fairly common in the Fells; this is the largest one at 32 inches (81.3 centimeters) DBH. It occurs in the newly designated Sugar Maple Middlesex Fells 3 NORMAN B. lEvENTHAl MAP CENTER AT THE BOSTON PuBlIC lIBRARy Middlesex Fells map from 1905. intentionally cultivated landscapes. As Elizur's daughter Ellen wrote, \"What we wanted in the Fells was a bit of nature so conveniently in our midst that we might watch its workings ... we wanted dark crowded places, even jungles ... marshes into which one might wade after reeds and bright berries, brooks where the border growth and waters frolic together ...\" A TrAcT oF WilD STony HillS The Fells is a dissected upland with narrow north 4 Arnoldia 70\/3 WAlTER KITTREDGE Middlesex Fells 5 MIKE RyAN Authority. The three reservoirs on the western side of the Fells were created out of the large Turkey Swamp before the reservation was set aside, and are owned (along with the adjacent land) by the town of Winchester. All of these reservoirs were included in both the original floristic survey of 1894 6 Arnoldia 70\/3 Middlesex Fells 7 rEcEnT FloriSTic SUrvEyS Deane's Flora reported a high level of biodiversity in the Fells, but a 1996 article by Drayton and Primack indicated an alarming loss of species. The article was based on a centennial study of part of the western Fells, done in the early 1990s by Brian Drayton for his master's degree. In the early 2000s, Bryan Hamlin began to question the validity of the 1996 report, after finding many of the \"missing\" plants in the area of study. As a result, he began a systematic resurvey of the entire Fells, assisted by Betty Wright, Don lubin, and others. At an NEBC meeting in 2006, Bryan Hamlin told me that he was working on a new flora of the Fells, and I agreed to help him with the difficult graminoid taxa--the grasses, rushes, and sedges. Over time, the current survey became a community effort among members of the NEBC, with a long list of local botanists contributing their expertise, very much like the original collaboration that led to the formation of the Club. For his survey, Drayton excluded ferns, graminoids, and aquatic taxa. Comparing the same set of plants from the same area, our survey found 564 taxa (355 native), while Drayton and Primack only reported 331 taxa (244 native) with a \"loss\" of 155 taxa since the Deane Flora. Our survey was able to find 105 of these reportedly lost taxa, 83 of them within their study area. The most likely reason for this large discrepancy was that Drayton's survey consisted mainly of a single person surveying for only 300 to 400 hours over three years, versus our team effort of about 2,000 hours over nine years. Drayton's work was also hindered because he wasn't allowed to collect specimens, which could lead to misidentifications. A large study of surveying techniques found errors of misidentification averaged about 5%, and that overlooking plants averaged 17%. After examining the Deane Flora vouchers, we found about a 4% error rate in misidentification. As stated in our Rhodora article, \"The level of expertise of the surveyors, the level of teamwork, and man-hours spent surveying all affect accuracy.\" For the 1890s survey, Manning defined four frequency categories--common, frequent, occasional, and rare--that the surveyors then reported according to their individual qualita- tive assessments. In order to create a quantitative measurement of frequency, we divided the Fells into eight approximately equal-sized sectors. Based on the number of sectors in which a plant was found, it was scored as common when found in seven to eight sectors; frequent in five to six; occasional in three to four; and rare in one to two. In order to obtain these data we conducted what were in effect eight minisurveys. Our examination of Deane's Flora and vouchers showed 680 vascular taxa (570 native, 110 non-native) for 1896, while our survey found 868 taxa (563 native, 305 non-native). This comparison of the two survey totals shows a tripling of non-native plants. While there was little net change in native plant numbers, there was a significant change in the composition. One hundred twenty-five native taxa that were reported in 1896 were not found by our survey, while we discovered or reconfirmed 119 BRyAN HAMlIN The parastic American squawroot was singled out by Deane as being the rarest plant in the Fells. It has since increased significantly in frequency, as we found it in five out of eight sectors. 8 Arnoldia 70\/3 Middlesex Fells 9 NANCy ROSE these were not previously known to occur in the Boston Basin Ecoregion, and two of them were newly designated during the course of this survey. One of these was Sugar Maple-OakHickory Forest, which is similar to Rich Mesic Forest, and only occurs on the south side of Bear Hill. The most prominent priority habitats were Rocky Summits, Pitch Pine Scrub Oak Communities, and vernal Pools. Over 100 vernal pools of varying size and hydrology have been identified, making the Fells a hotspot for vernal pools. The great diversity of habitats in the Fells can be accounted for by the diversity of geology and topography, in turn resulting in a high diversity of plants. Given the loss of land and the changing habitats over time, it is not surprising that there would be a significant change in the composition of the flora. The cessation of logging has allowed the forest to mature, with some areas starting to approach the characteristics found in old growth forest. Within the forest matrix, frequent anthropogenic fires continue to create a patchwork mosaic of different aged successional growth contributing to diversity. These burns have been kept small by the suppression of fires since the 1920s, which, along with increasing rainfall, has led to the overall favoring of mesophytic plants like beech and maple. During our survey a beaver dammed Whitmore Brook, creating a pond and marsh out of a red maple swamp, which resulted in an influx of new plants. Our survey found wetlands plants to be particularly opportunistic in responding to varying water levels and habitat succession. Studies of urban forests have found that the rarer plants with low population numbers are more susceptible to local extirpation. In the Fells about 60% of the taxa that were rare in 1896 are still extant, and almost half of those have increased in frequency. There are two state-listed rare species and eleven others which are watch-listed as potentially becoming rare in the state. Most of these rarer plants are herbaceous; woody plants are generally more abundant and more persistent. Some of the rare plants are ephemeral in nature, depending on successional habitats, and can come and go in a single season. We observed that orchids like Wild columbine (Aquilegia canadensis) is common in woodlands throughout the eastern United States and Canada, including Middlesex Fells. nodding ladies'-tresses (Spiranthes cernua) that were locally abundant in one year often went dormant and were very scarce the next. Other rare plants are restricted by only growing in habitats that are uncommon in the Fells. One of the factors that contributed to loss of plant populations was the replacement of the native oak forest around the Winchester Reservoirs with non-native evergreens. Another factor was construction. When Interstate 93 was built through the middle of the Fells in the late 1950s it destroyed a large area that included the only large fen habitat. A less obvious yet important negative factor is fragmentation caused by recreational overuse. There are 36 miles of fire roads and 75 miles of trails in the Fells, with a large proportion of these trails being created by users, resulting in very few large trailless areas. This extensive network of trails is an avenue for invasive plants to become widely established, evidenced by their abundance along the trails. 10 Arnoldia 70\/3 WAlTER KITTREDGE Middlesex Fells 11 beetles (Galerucella spp.) that eat purple loosestrife (Lythrum salicaria) has helped reverse the advance of this invasive plant, which was dominating wetlands. Although the deer population is relatively small, it still has had a negative effect on native lilies, which are also eaten by the non-native scarlet lily beetle (Lilioceris lilii). Other insects that pose potential future threats to the Fells forest include the emerald ash borer (Agrilus planipennis) and Asian longhorned beetle (Anoplophora glabripennis). Research in biological controls is ongoing and may eventually aid in controlling these highly destructive insects. Deane, W., ed. 1896. Flora of the Blue Hills, Middlesex Fells, Stony Brook, and Beaver Brook Reservations of the Metropolitan Parks Commission, Massachusetts. C. M. Barrows and Co., Boston, Massachusetts. Drayton, B. 1993. Changes in the flora of the Middlesex Fells, 1894"},{"has_event_date":0,"type":"arnoldia","title":"The Quest for the Hardy Southern Live Oak","article_sequence":2,"start_page":12,"end_page":24,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25547","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d24ea36b.jpg","volume":70,"issue_number":3,"year":2013,"series":null,"season":null,"authors":"Aiello, Anthony S.; Dosmann, Michael S.","article_content":"The Quest for the Hardy Southern Live Oak Michael S. Dosmann and Anthony S. Aiello CHrIS EvANS, IL WILDLIfE ACTION PLAN, BugWOOD.Org I t's no secret that individual plants within a species can vary in appearance--just peruse the range of Japanese maples (Acer palmatum) for sale at your local nursery. All belong to a single species, yet show diversity in traits like growth habit, foliage color, and leaf shape. It's also old news that individuals can vary according to provenance (geographic source); winter hardiness is frequently noted as one of those variable physiological traits. Although he was not the first to note this phenomenon, botanist and plant explorer Joseph Hooker provided an early description in1853. In an introductory essay preceding his notes on the flora of New Zealand, he described differences in the hardiness of Himalayan plants, \"depending upon the altitude at which they were gathered.\" Specifically, \"some of the seedling Pines whose parents grew at 12,000 feet appear hardy, Southern live oaks (Quercus virginiana) draped with Spanish moss line the road whilst those of the same species at Wormsloe, a historic colonial estate in Savannah, Georgia. from 10,000 are tender. The common scarlet Rhododendron of Nepal and the component of the missions of our respective North-west Himalaya is tender, but seedlings arboreta. When adding accessions, we want to of the same species from Sikkim, whose parents capture as much variation as possible within a grew at a greater elevation, have proved perspecies, so we often collect from multiple popufectly hardy.\" A few years ago, we wrote about lations within a species' range. This is standard C. S. Sargent's interest in acquiring cedar of practice for species in our core, or high-priority, Lebanon (Cedrus libani) germplasm that would collections that are already well adapted to our prove to be hardy in Boston (Aiello and Doslocal Arboretum conditions. However, for spemann 2007). He succeeded by obtaining seeds cies like C. libani that are not typically winter from Turkey, and those plants and others from hardy in our climate, we must seek specific that region have fared notably well in Philadelprovenances that may hold hardier populations. phia and Boston as well as colder climes, while One of those marginally hardy species that accessions from other provenances have failed. has evaded our grasp so far is the southern The cedar of Lebanon story points out the live oak (Quercus virginiana), whose massive, ongoing importance of plant exploration, a vital gnarled form--often draped in Spanish moss Southern Live Oak 13 (Tillandsia usneoides)--conjures up images of the antebellum South. This oak often exceeds 50 feet (15.2 meters) in height, but it is the spread that typically draws our attention. Almost always wider than tall, the colossal sweeping branches of old trees are a marvel. The common name \"live oak\" refers to the typically evergreen leaves, stiff and shiny on the top, and gray-tomentose on the bottom. However, during particularly cold spells the species may shed some of its leaves and is regarded as brevideciduous. Tolerant of drought as well as soil salinity and salt spray, southern live oak is often categorized as a \"tough plant,\" aside from winter hardiness issues. THe QueST BeginS Quercus virginiana has leathery, usually evergreen leaves. florida peninsula, turning northward to follow In 140 years of acquiring and testing species the coasts of georgia, South Carolina, North from all over the temperate world, the Arnold Carolina, and southern virginia. flint (1997) Arboretum has never even attempted to grow noted that while the species' useful range as Q. virginiana. That the Arboretum had tried-- a landscape plant is uSDA Zone 8b (average and failed--to establish hardy plants in the colannual minimum temperature 15 to 20 14 Arnoldia 70\/3 Southern Live Oak 15 the lab of Jeannine Cavender-Bares at the university of Minnesota has yielded interesting information on its ecology. Her lab found that Q. virginiana, like many other temperate species, varies in leaf and stem hardiness as a function of latitude: the more northern populations possess greater hardiness (Cavender-Bares 2007; Cavender-Bares et al. 2011; Koehler et al. 2012). In these studies, the lowest temperature that plants were exposed to (and survived) was 14 16 Madison university, 35 to 29). We commented that either the students were notably well behaved, or the landscape services department worked through the evening hours. using directions provided by Professor of Biology John Hayden, we were able to easily find the various specimens, many of which had been planted in the last few decades. Although we had seen the occasional Q. virginiana before, this site gave us our first chance to really observe the species in depth. Our first two collections were from trees growing near Westhampton Lake. The first tree, rounded and spreading in form, was about 15 feet (4.6 meters) tall and twice as wide; we estimated that it had been growing in that location for 10 to 15 years. And it was loaded with acorns, most with bright yellowish green nuts and tawny brown caps. However, a few had started to turn the typical mature color, a rich burgundybrown. The branches were dense, with short internodes, and thickly set with leathery, oblong to oval leaves. Considering their form and (brevi)evergreenness, we thought they would make great screens. As was our protocol for the entire trip, we gathered germplasm in the form of acorns, made herbarium vouchers from cut twigs (complete with the acorns), and of course jotted down copious collection details that pertained to the trees as well as the local conditions and environment. The second collection was from a nearby tree, smaller and younger than the first, but similar to another six growing nearby. undoubtedly the campus was trying to establish a grove of these trees in this area. Before leaving the university, we located and collected from two trees, older than the first, which were growing near a dining hall. changes in Plant Hardiness Zones IN JANuAry 2012, the united States Department of Agriculture unveiled its new Plant Hardiness Zone Map (PHZM) (http:\/\/planthardiness.ars.usda.gov\/PHZMWeb\/), a development that was long anticipated by gardeners and researchers. Like its earlier incarnations, the new PHZM provides guidelines to predict a region's average annual minimum temperature, a vital statistic in determining whether or not a plant may survive the winter in a particular area. Last updated in 1990, the map now features a number of significant features. for one, it has gained interactivity through a geographic Information System (gIS) that enables users to zoom in at regional and state levels; it also has a tool to identify a zone by zip code. Data quantity and quality represent marked improvement in the map's reliability-- the new PHZM utilizes 30 years (1976 allow gardeners to reliably grow Stachyurus praecox and Chimonanthus praecox, which are currently hardy only in protected microclimates. And, if we are lucky, Philadelphia and Boston can add Quercus virginiana to that list. Our next destination--after an amazing lunch at Buz and Ned's BBQ--was Bryan Park, a historic richmond landscape founded in 1910. We expected to find small, rounded trees similar to those we had found at the university earlier in the morning. However, what we did find were three very large individuals, just down the hill from the gatekeeper's House on the park's northeast side. Heights ranged from 30 to 40 feet (9.1 to 12.2 meters); each was rounded, usually twice as wide as tall, and with gnarled, twisting stems and branches. Only two of the trees (with dbh values of 35 and 39 inches [89 and 99 centimeters], respectively) bore acorns. Although we do not have any records to confirm this, based on their size we assume that the trees date back to the founding of Bryan Park and approach the 100 year mark. If so, they certainly would have survived the frigid winter of 1940. TO WiLLiamSBuRg We departed richmond in the early morning of October 22nd, and by 9:00 a.m. arrived at our next destination: the College of William and Mary in Williamsburg. Beth Chambers, curator of William and Mary's herbarium, was a great help to our efforts. Prior to our arrival, 18 Arnoldia 70\/3 Southern Live Oak 19 A grove of old southern live oaks at the edge of the Parade Ground of Fort Monroe. The moat surrounding Fort Monroe contributed to its defenses; mature southern live oaks can be seen growing within the fort's interior, above and to the right of the casement. resources Management at the fort. We were thankful for the lead. Perched at the ocean's edge, the fort has a rich history that dates to the early seventeenth century. It had been occupied by the military until its recent decommissioning in 2011, and it is now a National Monument. The massive six-sided stone structure is the largest of its kind in North America: 63 acres of land surrounded by walls and an impressive moat. Construction of the current fort took 15 years to complete and the final phase (finished in 1843) was overseen by robert E. Lee. In an ironic twist, such was its fortitude that it was never lost to the Confederacy. We arrived in the late afternoon of the 22nd to meet Eola, who enthusiastically showed us around the facility and explained some of its fascinating history. We also returned on the morning of the 24th to visit with her, as well as Joshua gillespie and robert Kelly of the fort Monroe Authority. Inside the buttressed edifice we found a composite of former army barracks, period officer quarters, office and training facilities, storage buildings, a chapel, and a museum, as well as nearly 350 southern live oak specimens scattered throughout. Perhaps the most impressive is a large grove that grows along the south and west edge of the interior parade ground. Some trees stood as lone sentries, while others grew in small groups, sometimes arching over the sidewalks and defying gravity. Most were no taller than 35 to 40 feet (10.7 to 12.2 meters), and all had dramatic, ethereal forms, the result of decades and even centuries of difficult environmental conditions including drought, intense heat, and salt spray (even inside the fort's walls). No doubt, the grandest of these was the Algernourne Oak, a leviathan estimated to be over 450 years old. This tree has a basal diameter of 90 inches (228.6 centimeters), with two massive leaders diverging about 3 20 Arnoldia 70\/3 Southern Live Oak 21 in the parade ground, we imagined ourselves dressed in full uniform, performing drills and marching for hours under the hot sun and dry, salty breeze--those trees would be considered sacred! The trees were in remarkably good condition considering their age, size, and the heavy impact of human activities on the site. Many of them showed the marks of time but they were mostly healthy and growing well, a testament to the resilience of southern live oaks. fiRST Landing We dedicated the 23rd to surveying the flora of first Landing State Park, which lies on Cape Henry between Norfolk and virginia Beach. Its current name, changed from Seashore State Park in Tony Aiello stands near a cluster of Osmanthus americanus along the 1997, acknowledges this site as the namesake Osmanthus Trail at First Landing State Park. location where the virginia Company first landed in 1607 prior to settling Jamesfinding them was quite easy thanks to town. The park covers about 3,000 acres, and our earlier planning conversations with Erik comprises eight upland plant community types Molleen of the virginia Department of Conthat range from dune crests to mesic forests servation and recreation; the fact that there (Clampitt 1991). Our initial foray was into the was an Osmanthus Trail in the park was also mesic forests where several of our non-oak helpful. Osmanthus americanus specimens collecting targets were to be found: devilwood were numerous and scattered throughout the (Osmanthus americanus) and swamp bay (Perunderstory. They became easy to identify from sea palustris). Like southern live oak, these two a distance because their glossy green leaves are species of shrubs or small trees are near or at arranged oppositely, as with other members their northernmost ranges in virginia. And, for of the olive family (Oleaceae). At the Arnold reasons similar to our quest for hardy southern Arboretum, this species has proven to be quite a live oak germplasm, we were anxious to locate challenge to cultivate because of cold hardiness and collect from these species. issues. One clone, a cultivated lineage from Spring grove Cemetery in Cincinnati, Ohio, has been reliably hardy in Boston. Likewise a plant at the Morris Arboretum has survived but not thrived since it was received from a local nursery in 1962. Wild-provenance material has long been a target because of the species' botanical and ornamental appeal. Its broadleaved evergreen foliage provides winter interest, and the small, creamy white flowers in spring are a delight to the nose; their mellic scent beckons from great distances. We were able to collect fruits--bright green drupes at this stage--from many trees in the woodland. Persea palustris also dotted the understory, Devilwood (Osmanthus americanus) bears sweetly fragrant flowers in the spring. and, like devilwood, has large, elliptic, evergreen MIKE HOgAN, HTTPS:\/\/fP.AuBurN.EDu\/ SfWS\/SAMuELSON\/DENDrOLOgy 22 Arnoldia 70\/3 Southern Live Oak 23 The water table was down considerably at First Landing State Park, exposing the buttressed trunks and knees of the bald cypress (Taxodium distichum). branching form. One of the larger trees we found had three stems measuring 12.5, 17, and 21 inches (31.8, 43.2, and 53.3 centimeters) in diameter at 12 inches (30.5 centimeters) above the ground. Despite the stressful environment, trees were healthy and there was noticeable regeneration of young seedlings in the understory, which is always a good sign. rather than focus on individual trees at this site, we maximized the amount of genetic variation in the collection by gathering acorns from 12 trees. Some trees were so fecund and at perfect ripeness that we could easily shake the branch and scores of the nuts would drop from their caps. nexT STePS Although the fieldwork is complete, the data are in the databases, and the herbarium specimens are mounted, much work remains ahead of us. Each of our institutions is hard at work germinating the seeds from the various collections made on the trip--twelve separate Q. virginiana collections, plus one each of the Persea, Osmanthus, and Q. 24 Arnoldia 70\/3 "},{"has_event_date":0,"type":"arnoldia","title":"Mark Catesby: Pioneering Naturalist, Artist, and Horticulturist","article_sequence":3,"start_page":25,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25545","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270896d.jpg","volume":70,"issue_number":3,"year":2013,"series":null,"season":null,"authors":"Yih, David","article_content":"Mark Catesby: Pioneering Naturalist, Artist, and Horticulturist David Yih T his April marks the 301st anniversary of naturalist Mark Catesby's arrival in Williamsburg, Virginia, to begin the first of two exploratory sojourns he would make in the American colonies. A dabbler in watercolors from a family of provincial English lawyers, Catesby was twenty-nine when he stepped off the ship to begin the adventure that would determine the course of his life and culminate in his monumental work on North American flora and fauna, The Natural History of Carolina, Florida, and the Bahama Islands. The lavishly illustrated work would be hailed in the Philosophical Transactions of the Royal Society of London as \"the most magnificent work ... since the art of printing has been discovered\" (Mortimer 1748). It would stand as a benchmark in American natural history throughout the eighteenth century and be deemed \"the most splendid of its kind that England had ever produced\" (Pulteney 1790). Though little documentation of Catesby's early life exists, it is generally supposed that his interest in the natural world had been stimulated by his uncle Nicholas Jekyll, an avid gardener who introduced the young man to John Ray, \"the foremost English naturalist of the late seventeenth century ... whose systems would dominate English natural history until the adoption of Linnaean classification\" (Frick 1974). The best glimpse into Catesby's preoccupations as he first arrived in America to visit his sister's family and have a look around comes in his own words: \"... my Curiosity was such, that not being content with contemplating the Products of our own Country, I soon imbibed a passionate Desire of viewing as well the Animal as Vegetable Productions in their Native Countries; which were Strangers to England. Virginia was the Place (I having Relations there) suited most with my Convenience to go to, where I arriv'd the 23d. of April 1712. I thought then so little of prosecuting a Design of the Nature of this Work, that in the Seven Years I resided in that Country, (I am ashamed to own it) I chiefly gratified my Inclination in observing and admiring the various Productions of those Countries, only sending from thence some dried Specimens of Plants and some of the most Specious of them in Tubs of Earth, at the Request of some curious Friends ...\" (Catesby 1731) Perhaps Catesby could afford to be a bit modest by the time he wrote these prefatory words of his celebrated magnum opus. In reality, when he returned to England after seven years in the colonies, he \"brought with him an extensive knowledge of New World flora and fauna as well as an impressive cache of drawings of animals and plants never before seen by English naturalists\" (Meyers and Pritchard 1998). These were sufficient to attract the interest of the eminent English botanist William Sherard, who happened to be in the process of organizing sponsors to send a naturalist across the Atlantic to explore and document the living wonders of America, especially those that might have scientific, economic, ornamental, or curative value. Whom to send on this mission was an issue yet to be resolved. But an ability to render accurate images of the new finds would be a significant qualification. Impressed by Catesby's work, Sherard wrote to an acquaintance, \"He designs and paints in water colours to perfection.\" Catesby got the job, and with the support of a dozen backers--including a number of aristocrats as well as the President and several members of the Royal Society--set out on his second journey, arriving in Charleston, South Carolina, in 1722. With the funds and trust that were now invested in him, he threw himself into 26 Arnoldia 70\/3 Mark Catesby 27 his work, resolving never to visit the same area twice during the same season. The frequent clamoring of his impatient backers for specimens sometimes hampered his efforts at what he saw as the main thing to be accomplished: an illustrated record of the plants and wildlife of America. But he persevered and for four years ranged from coastal plains to Appalachians and from the Carolinas south through georgia, Florida, and the Bahamas, collecting, documenting, and painting as he went. Upon his return to England in 1726, Catesby took a job as a nursery horticulturist and began work on the great book he envisioned. The project would take more than twenty years to complete. And he would have to publish it himself. In a practice common at the time, Catesby solicited subscribers by issuing a prospectus describing the proposed publication and his qualifications for undertaking it. Subscribers would make advance payments, and these would help defray the costs of producing the books. Catesby gave persuasive evidence of the worthiness of his project by listing in the prospectus the names of the twelve eminent men who had sponsored his second trip and by publicly exhibiting the drawings and watercolors he had brought with him from the colonies. Ultimately, 155 persons and institutions signed on, enough to set the project in motion. In order for the illustrations to be printed, they would have to be engraved into copper The Blue Bird and Smilax non spinosa, humilis Eastern bluebird (Sialia sialis) and sarsaparilla vine (Smilax pumila) 28 Arnoldia 70\/3 Mark Catesby 29 plates. Catesby had hoped to have the work done by the expert engravers of Amsterdam or Paris, but given the number of plates involved--220 would grace the finished work-- the expense proved prohibitive. Undeterred, he studied the technique of etching with Joseph goupy, a French printmaker and art instructor then living in England, and proceeded to etch all of the plates himself. He published the work in installments of twenty plates with accompanying bilingual English 30 Arnoldia 70\/3 Mark Catesby 31 The Painted Finch and the Loblolly Tree Painted bunting (Passerina ciris) and loblolly bay (Gordonia lasianthus) 32 Arnoldia 70\/3 Mark Catesby 33 the direction of twist shown in the twining of Catesby's sweet potato plant (Ipomoea batatas) is incorrect. Catesby himself recognized that his artistic skills were limited by his lack of expertise in perspective but felt that his flat depictions were sufficient for the purpose of delineating species. In time, his work was superseded by the achievements of later generations, and Catesby's renown faded. \"After the American Revolution, interest in Catesby's work, as with most things American, waned in England. And as the scientific community became increasingly specialized, . . . Catesby's generalist approach fell into disfavor. By the time John James Audubon set off to paint in South Carolina nearly a century later, Catesby had been almost forgotten.\" (Amacker) In recent decades, however, a new appreciation of Catesby's contribution has emerged. With the perspective of two-and-a-half centuries, it has become clear that Catesby's work was innova- The Blueish Green Snake and Frutex baccifer, verticillatus tive and ahead of its time. He Rough green snake (Opheodrys aestivus) and American beautyberry (Callicarpa americana) broke from the stilted bird profiles typical of the times to include dynamic logical characteristics. In the case of birds, he images of birds in motion. The bald eagle in often commented on aspects of nest-building, full swoop, bearing down upon its prey in the feeding, and migratory behaviors. He authored very first plate is an example. He was the first the first scientific paper (Catesby 1746-7) to to depict birds against botanical backgrounds. accurately address the phenomenon of bird More importantly, in choosing these backmigration (earlier theories had birds hibernatgrounds, he made a conscious effort to depict ing in caves or under water during the winter ecological relationships, frequently showing months). For these reasons, and in considerbirds with the plants on which they feed or in ation of the many new bird species he brought which they nest. His texts go beyond describto light, Catesby has been called the founder of ing morphology to reveal behavioral and ecoAmerican ornithology (Frick 1974). 34 Arnoldia 70\/3 Mark Catesby 35 Steuartia Silky camellia (Stewartia malacodendron) [Ed. note: Though he named the genus in honor of John Stuart, Linnaeus spelled it as Stewartia. This is still the generally accepted spelling, though some taxonomists spell it as Stuartia. Catesby's spelling seems to split the difference.] 36 Arnoldia 70\/3 "},{"has_event_date":0,"type":"arnoldia","title":"Book Review: Knowing Nature: Art and Science in Philadelphia, 1740-1840","article_sequence":4,"start_page":37,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25544","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d2708928.jpg","volume":70,"issue_number":3,"year":2013,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Book Review: Knowing Nature: Art and Science in Philadelphia, 1740 38 Arnoldia 70\/3 Book Review 39 story of John Bartram's involvement in the discovery of ginseng in Pennsylvania (1739) and his efforts to collect plants for his patron, Peter Collinson, who was interested in establishing a business exporting American ginseng from England to China. While I have read about this story before, the six beautiful images of ginseng (including a botanical specimen collected by Bartram) that illustrate Janice Neri's chapter on the China trade give this version a vitality that text alone does not provide. Mark Laird's chapter on \"The American Connection in Georgian Pleasure Grounds\" traces how the interest in and importation of North American plants and animals into England changed the nature of designed English landscapes. In a similar vein, I found Lisa Ford's chapter about Fran"},{"has_event_date":0,"type":"arnoldia","title":"Betula dahurica: A Special Birch Tree","article_sequence":5,"start_page":40,"end_page":40,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25543","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270856f.jpg","volume":70,"issue_number":3,"year":2013,"series":null,"season":null,"authors":"McAllister, Hugh","article_content":"Betula dahurica: A Special Birch Tree Hugh McAllister I n the United Kingdom, Betula dahurica has a reputation for not making a well-shaped tree, as it often suffers repeated dieback and poor growth because of late spring frosts and inadequate summer heat. One specimen in the Arnold Arboretum (overhanging the road on Bussey Hill) shows the typical \"witches' broom\" growths caused by such repeated dieback, but most trees of B. dahurica in the Arboretum have made good specimens. Particularly noteworthy is a tree of Japanese origin (accession 1015-80-A) just off Conifer Path near the bamboo collection. Dahurian birch is noted for its peeling, papery bark (similar to river birch, B. nigra) and this specimen has particularly attractive shaggy curls that have a redder color on their inner surface than some other Arboretum specimens. The color of the inner surface contrasts nicely with the creamy white of the outer surface of the curls and the unpeeled sections of bark on the branches. Betula dahurica is native to China, Japan, Korea, eastern Mongolia, and far eastern Russia. Accession 1015-80-A is of special interest since B. dahurica is endangered in Japan, being known primarily from a small population near Nobeyama in Nagano Prefecture in the central part of the main island of Honshu (where this accession was collected). There is another small population in the northern island of Hokkaido and one on Iturup in the Kurile Islands, which were Japanese before being occupied by Russia at the end of World War II. Of genetic interest, these offshore island populations are hexaploid (6 times the base number for birches of x=14) with a chromosome number of 2n=84, whereas the extensive populations on the Asiatic mainland all appear to be octoploid with 2n=112. This means that the island populations are unlikely to interbreed freely with the mainland populations, are genetically distinct, and, if they can be recognized by their appearance, should be named as a distinct species. Three cuttings from the tree in the Arboretum have been rooted and are now growing in the nursery. The only other known trees from the Nobeyama provenance in cultivation are a single tree at Dawyck, a satellite garden of the Royal Botanic Garden Edinburgh in southern Scotland, and six trees at Ness Gardens, the University of Liverpool Botanic Gardens near Chester in northwest England. Trees from this provenance grow far better in the United Kingdom than any from continental Asia, presumably because of the greater similarity of our climate to the maritime climate of Japan. Since the Nobeyama trees are genetically distinct and rare in the wild, they are clearly of conservation significance and efforts should be made to have breeding populations for seed production in cultivation. Most species of birch are self-incompatible (self-sterile), so at least two different seedling trees are needed for seed production. Fortunately we have this at Ness and, despite the large number of other birch species in the surrounding garden, seedlings from the cultivated trees seem to be mostly coming true (i.e., are not hybrids with other species). Accession 1015-80-A is producing some viable seeds, so it will be interesting to sow this and see what the seedlings are. If the parent tree is totally self-incompatible then all the seedlings will be hybrids. No known hybrids of B. dahurica have ever been reported, and certainly no hybrids of the Nobeyama provenance, so, if we can identify what the other parent(s) might have been, it will tell us what other species B. dahurica can hybridize with. Any such hybrids could be of horticultural interest since B. dahurica may be resistant to bronze birch borer. Alternatively, accession 1015-80-A could have a limited degree of self-compatibility (resulting in a low percentage of viable seeds) and at least some of the seedlings could be the result of self-fertilization. This could result in some dwarf or other abnormal growth forms as a result of inbreeding depression--this is the probable mode of origin of many dwarf conifers. No doubt this species, and the Japanese provenance in particular, will continue to be studied, conserved, and propagated at the Arnold Arboretum, Ness Gardens, and other botanical institutions. Hugh McAllister is an honorary lecturer at the University of Liverpool and was recently a Sargent Award visiting scholar at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23431","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160896e.jpg","title":"2013-70-3","volume":70,"issue_number":3,"year":2013,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Mystery of Seasonal Color Change","article_sequence":1,"start_page":2,"end_page":10,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25542","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270816b.jpg","volume":70,"issue_number":2,"year":2012,"series":null,"season":null,"authors":"Lee, David","article_content":"The Mystery of Seasonal Color Change David Lee ... the gods are growing old; The stars are singing Golden hair to gray Green leaf to yellow leaf,--or chlorophyll To xanthophyll, to be more scientific ... Edwin Arlington Robinson (Captain Craig) T hroughout New England each autumn-- early October in some parts and as much as three weeks later in others--the pageant of color change in our forests unfolds. Though less noticed, in the springtime these forest canopies take on delicate pastel colors as buds swell and leaves expand. In the last 15 years, our understanding of the science behind color change has begun to emerge, with two different but not mutually exclusive hypotheses being formulated and defended. I have been involved in the research and debate on these color changes, and why that is so is a bit of a mystery in itself. After all, I grew up on the cold desert of the Columbia Plateau in Washington State, where the predominant colors were the grays of sagebrush and other pubescent shrubs. Occasionally I visited the forests of the Cascade Range to the west, witnessing the dark greens of conifers, occasional yellows of cottonwood, birch, and willow in the autumn, with just a few splashes of the reds of the Douglas maple (Acer glabrum var. douglasii). I did enjoy the autumn colors of the mid-Atlantic and Midwest forests as a graduate student and post-doctoral fellow, but was too busy in the laboratory to think much about that color. Then I moved to tropical Asia--Malaysia specifically--and took notice of the differences in tropical rainforest vegetation, which I have been studying ever since. I was particularly struck by the red colors of leaves, both on the undersurfaces of understory plants and the expanding leaves of giant trees (so colorful that from a distance they looked to be in flower). Yet, few of A red maple leaf shows developing red autumn color along with still-green sections. This micrograph of a red maple (Acer rubrum) leaf shows that it contains both red anthocyanins and yellow xanthophylls. Seasonal Color Change 3 the leaves turned red before falling from trees, and the canopies remained green because leaf fall was staggered. I began studying that red color--which led me back to the autumn reds of New England forests. A PAlETTE of PigMEnTS As poet Edwin Arlington Robinson partly described, the colors of leaves are the products of pigments produced in their internal tissues. Chlorophylls produce greens, xanthophylls produce yellows and oranges, and anthocyanins (left out by Robinson) produce reds. The leaf tissue is like the thick paper employed in watercolor painting. If you consider that most of the leaf consists of cellulose fibers, the similarities are particularly strong. The interior leaf volume, with its numerous air chambers facilitating the exchange of gases that supports photosynthesis, strongly scatters light, allowing some to reflect and some to be transmitted through the leaf. The leaf pigments are then All IMAgES bY ThE AUThOR like the soluble pigments in watercolors, and color is produced subtractively. Chlorophyll produces a green color because it absorbs light in the blue and red wavelengths. Xanthophylls produce yellow because they absorb blue into green, and anthocyanins red because they absorb even more green into blue. These pigments can combine to produce oranges (yellow and red), or even brown (green and red). The colors of spring and autumn are produced by these pigment combinations in leaves. When I was a college student of botany in the 1960s, the textbooks taught us that autumn colors were produced by the loss of chlorophyll unmasking the yellows of xanthophyll and reds of anthocyanin pigments, and that the colors had no function. To me, this did not seem right for the reds of anthocyanins, because I knew that these pigments are quickly synthesized in leaves. When I began to work at Florida International University in Miami in 1980, I turned my attention to the young leaves of mango and A red maple glows in autumn color at the edge of Connor Pond, between the towns of Petersham and Barre in central Massachusetts. 4 Arnoldia 70\/2 Seasonal Color Change 5 entire tree for the following year, and we speculated that the advantage could be to protect the leaves during the process of the breakdown of chlorophyll. leaves carefully disassemble the chlorophyll and associated proteins during senescence, and much of the nitrogen-containing compounds are resorbed in the woody tissues for use the following spring. We discussed a plan of action, and Missy supported my application for a bullard Fellowship at the harvard Forest in Petersham, Massachusetts, which I used from late summer through early winter in 1998 and 2004. During those sojourns in central Massachusetts, with frequent cultural trips to Cambridge and boston, I observed the changes in the forest pretty much every day, and made observations and physiological measurements of leaves during the process of leaf senescence and color change. When the snow started falling in early December, I high-tailed it back to balmy Miami. Missy and I collaborated with her Ph.D. student Taylor Feild (now at the University of Tennessee) and harvard Forest scientist John O'Keefe, who had been observing the phenology (when trees leaf out, when they flower and fruit, and when the leaves change color and fall from the trees) of common tree species at the forest starting in 1991. We found that most of the trees and shrubs (62 of 89, or 70%) produced anthocyanins during senescence, starting when leaves had already lost about half of their chlorophyll. Such leaves appeared red, red-orange, bronze, and even brown in color. The precise colors depended on the mixtures of anthocyanins, chlorophylls, and xanthophylls. We studied the changes in pigment composition and physiology in leaves of individuals of 16 species, 8 with anthocyanins and 8 whose leaves turned yellow (they had residual xanthophylls but no anthocyanins). We found that anthocyanin concentration during senescence was correlated with lower nitrogen content, consistent with the prediction that more nitrogen could be resorbed by the woody Autumn leaf color in sugar maple (Acer saccharum) varies between (and even within) individual specimens. 6 Arnoldia 70\/2 Seasonal Color Change 7 That created (1) some good will, (2) the understanding that we didn't know much about either hypothesis (which is often the fuel for disagreement and animosity), (3) the most exhaustive review on the subject (see the citation at the end of this article), and (4) an exhortation that we should produce the missing data that would more critically test the hypotheses. because it involves animals, the co-evolution hypothesis has attracted the most research. When we observe color, we automatically think of signaling--as from attractive flowers to pollinators, and from colorful fruits to dispersers. Conversely, in the physiological hypothesis, color is merely the by-product of protective absorption at specific wavelengths we can't see. We now know that color, particularly yellow against a green background, repels visits by aphids, at least in the few trees that have been examined, especially European birch (Betula pendula). The evidence for red leaves is more controversial. Although there is some evidence of aphids avoiding red leaves, these insects seem not to have visual receptive cells sensitive in the red wavelengths. limited evidence supports the contention that repelling aphids reduces egg laying, decreases activity of these sucking insects the following year, and increases seed production. Marco's strongest supporting evidence is from wild apples, where autumn aphids lay more eggs on green rather than red leaves. Inside and outside views of autumn leaf color in red oak (Quercus rubra). Another weakness is a lack of evidence of reduced palatability, greater duEling hyPoThESES toxicity, or less nutrition in yellow or red Publications supporting the two hypotheses leaves, although such leaves would likely be stimulated additional research, along with a more advanced in senescence and thus less number of speculative reviews. A little aninutritious than green leaves. A model of the mosity arose between proponents of these kind of research needed was published in 2011 two views, even though the hypotheses were by Kevin gould and colleagues in a New Zeanot mutually exclusive. To Marco's credit, he land tree, horopito (Pseudowintera colorata), convened a meeting at Oxford in March 2008, that has red leaf margins. They found a toxic with researchers from both \"camps\" present. plant molecule, polygodial, was associated with 8 Arnoldia 70\/2 Seasonal Color Change 9 Though more subtle than in autumn, reddish color is also common in spring on expanding leaves, as seen here in woods near Beaverkill, New York. cence, or the pressures of herbivory. Many maples and oaks produce red colors during the autumn, and birches and beeches produce yellows. hoch has shown that the birches resorb nitrogen as well as the species with red anthocyanins, so there are likely to be other protective mechanisms that have evolved. The color production in geographically distant forests also varies greatly. Among the most spectacular color displays are autumn forest scenes in New England, and also in the southern Appalachians (residents of the great Smoky Mountains region think their colors are the most beautiful!). however, European forests produce little red, and a lot of yellow. Although red and yellow colors are produced in virtually all deciduous trees in New England, in other forests leaves may stay green. Isreali botanist Simca lev Yadun has speculated that variation in color production may be the result of the different tree species in particular forests, and their evolutionary histories. Past and future climates may affect color production by determining the distributions of different trees in forests. Diseases may also remove species and change the colors during autumn. For New England forests, I expect that a century ago there was much more yellow in the autumn color palette. The rapid decline of the American chestnut from chestnut blight 10 Arnoldia 70\/2 "},{"has_event_date":0,"type":"arnoldia","title":"Inside Plants: An Engineer's View of the Arnold Arboretum","article_sequence":2,"start_page":11,"end_page":19,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25540","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270bb6d.jpg","volume":70,"issue_number":2,"year":2012,"series":null,"season":null,"authors":"Gibson, Lorna J.","article_content":"inside Plants: An engineer's View of the Arnold Arboretum Lorna J. Gibson G ardeners tend their plants to produce a beautiful display in the garden or to harvest fruits and vegetables. Botanists study the anatomy, life cycles, and evolution of plants. Engineers, too, are interested in plants, although from a different perspective. Historically, engineers have been interested primarily in wood, because of its widespread use in everything from furniture to boats to buildings. But more recently, engineers have recognized that plants are very effective at resisting the loads they are subjected to (for instance, from the wind or from their own weight). Today, engineers study plants to learn what features make them so effective mechanically, with a view towards \"bio-inspired design\" of engineering materials and structures that exploit these features. In this article, I will take you for a walk through the Arboretum and describe a variety of plants and how they work from an engineering perspective. The Tour Begins Across from the Hunnewell Visitor Center, east of Meadow Road, lies the Cattails (Typha spp.) growing in the Meadow at the Arnold Arboretum. Meadow, a marshy area largely filled with cattails (Typha spp.). The leaves stand along their length; if you draw your thumbnail close to vertical and reach an impressive height, across the width of the leaf you can feel the often over 6 feet tall. As you walk past, you can ridges of the fibers. How do the long, thin leaves see the leaves bend in the wind; occasionally, stand up so tall? a sparrow or red-winged black bird lands on A look at the cross section of a cattail leaf the stem or its fuzzy, cylindrical seed head and reveals the answer. The cross section shows bends that over, too. If you look at the leaves up two outer faces connected by a number of ribs. close, you can see that they have fibers running At the very outer top and bottom surfaces, you RoBERT MAyER 12 Arnoldia 70\/2 Inside Plants 13 nAnCy RosE and grasses. A cross sectional view of an iris leaf shows that it has large dense fibers (called sclerenchyma) at the outer surface and a thick inner layer of foamlike cells (called parenchyma). When the leaf is bent, the dense fibers carry most of the high internal loads at the outside of the leaf. The separation of the denser, stiffer fibers by the inner foamlike layer increases the resistance of the iris leaf to bending. Engineers make use of the same concept (a \"sandwich structure\") in the design of downhill skis, lightweight panels for aircraft, and the blades of windmills, which often have two outer skins of carbon-fiber-reinforced plastic separated by a foam (or sometimes an engineering honeycomb) core. suPPorTing AcT If we walk back towards the Arborway Gate and look along Willow Path, we see the huge leaves of the butterbur (Petasites japonicus). How does the stem support such large leaves without falling over? The stem bends under the weight of the leaf and from wind acting on the leaf. The stem is roughly circular in cross secLoRnA J. GIBson The swordlike leaves of bearded iris have a \"sandwich structure\" that increases their resistance to bending. Don GALLER, MIT Dense outer fibers (schlerenchyma) are separated by foamlike parenchyma in this iris leaf cross section. An empty tube kinks when bent, but the foam-filled tube resists kinking. 14 Arnoldia 70\/2 nAnCy RosE Birds of a Feather Bird FeATher quills have a similar structure to iris leaves. As the bird's wings beat in flight, the feather is bent up and down, so that the bending loads are highest on the top and bottom of the quill. A sandwich structure, with dense faces at the top and bottom of the quill, would seem to be an ideal option. But bird feathers also have to resist twisting, or torsion, and the foam-filled closed tubes are adept at this. (You can try this simple experiment: take a drinking straw and twist it. now cut a slit along the length of the straw and twist it again. The straw is much better at resisting twisting when the cross section is closed and intact.) As with the butterbur, the foamlike core also helps resist kinking failure of the dense outer layer of material in the feather quill. CouRTEsy of AL REID Don GALLER, MIT Blue jay (Cyanocitta cristata). Cross section of a blue jay feather. bark of the cork oak is removed, it regrows, allowing harvesting of cork every 10 to 15 years. The cork cells are like little bellows: they are roughly box-shaped, but with corrugations running in one direction. When you compress the cork in the direction of the corrugations, they simply fold up, like a bellows, so that they do not expand in the lateral direction. This feature of cork is one reason cork works well at stoppering bottles. A rubber stopper, on the other hand, bulges out laterally when compressed, making it difficult to press into a bottle; for this reason, rubber bottle stoppers are always tapered. Trees = Wood When we think of the Arnold Arboretum, we think of trees. And when engineers think of trees, they inevitably think of wood. Wood is one of the structural materials used for the longest time in human history and is still one of the most widely used. The oldest known wooden boat is Cheops's 4,600 year old barge, found dismantled in a pit next to the Great Pyramid in Egypt. In the late 1600s, eastern white pines (Pinus strobus) from new England were a strategic resource for the British Royal navy. The tall, straight trunks of the pines were used as masts for ships; the taller the mast, the more sail area, the larger the ship, and the more cannons it could carry. And most houses in north America are still wood framed. north American woods are divided into hardwoods (deciduous trees that drop their broad leaves annually) and softwoods (conifers with needles that are typically, but not always, evergreen). While hardwoods tend to be denser and harder than softwoods, that is not always the case: for example, Douglas fir, a softwood, 16 Arnoldia 70\/2 LoRnA J. GIBson Inside Plants 17 Vessel t Oak wood, cross section and longitudinal section. t Tracheid t Fiber ray t t ray Cedar wood, cross section and longitudinal views. wood, this is equivalent to the fraction of the volume that is solid. In contrast, when a model honeycomb is loaded across the cells, it is much easier to deform the honeycomb, as the cell walls bend. Wood cells loaded across the grain also bend in a manner similar to the honeycomb; this can be seen most easily in a low density wood like balsa (Ochroma pyramidale) (see upper right images on page 18). If you take a ruler and bend it, it deforms much more than if you rest one end on a table and compress it from the opposite end with the same load. It is also less strong when bent: it is much easier to break the ruler in bending than by compressing it on end. We have already seen how, in a bent beam, the amount that the material stretches or compresses increases as the distance up or down from the middle of the beam increases: the thickness of a beam plays a greater role in resisting deflection or internal loads than the width. When loaded across the grain, the wood cell walls bend, giving much lower stiffness and strength across the grain than along the grain. This effect can be analyzed in more detail to show that the stiffness of woods loaded across the grain depends on the cube of the volume fraction of solid, and the strength (loaded across the grain) depends on the square of the volume fraction of solid. This leads to the great difference in the stiffness and strength in woods when loaded along and across the grain, a difference that is greater in lower density woods, such as pine, than in high density woods, such as oak. for instance, in Eastern white pine the compressive strength 18 Arnoldia 70\/2 Inside Plants 19 nAnCy RosE typically loaded in bending from the wind--the increase in denser material towards the outside of the cross section increases the resistance of the culm to bending deflections and loads, compared with a section with the same amount of material evenly distributed across the section. on our walk through the Arboretum, we have seen a variety of plants with different internal structures. Plants are often mechanically efficient, using material to resist internal loads where they are greatest. Engineers studying the mechanical behavior of plants take inspiration from them for the design of engineering materials and structures. Acknowledgements Bamboo sEM figure is from Gibson, L. J., M. f. Ashby, G. n. Karam, u. Wegst, and H. R. shercliff. 1995. The mechanical properties of natural materials II: microstructures for mechanical efficiency. Proceedings of the Royal Society of London A450: 141"},{"has_event_date":0,"type":"arnoldia","title":"Fruits and Nuts of the Villa Farnesina","article_sequence":3,"start_page":20,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25684","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e0d260af26.jpg","volume":70,"issue_number":2,"year":2012,"series":null,"season":null,"authors":"Janick, Jules","article_content":"Fruits and Nuts of the Villa Farnesina Jules Janick The loggia of Psyche in the Villa Farnesina. The paintings on the ceiling depict scenes from the love story of Cupid and Psyche. I n 1505, an extremely wealthy Sienese banker named Agostino Chigi (1466 The Villa Farnesina 21 Richly detailed festoons painted by Giovanni Martini da Udine contain thousands of botanical images. the decorations of his home, Chigi was more interested in the sensuality of the pagan world than on the artistic representation of suffering, torture, and death of the medieval Christian tradition. The decorations--painted between 1515 and 1517--involve love and marriage, perhaps in anticipation of Chigi's marriage to his longtime mistress in 1519, one year before his death. The loggia is presented as a tentlike pergola with images of two large tapestries painted on the ceiling as a roof, depicting a scene of the Council and the Banquet of the Gods--the climax of the Cupid and Psyche story. The arches of the ceiling are divided into spandrels that contain scenes of the heavenly adventures of Venus, Cupid, and Psyche, alternating with severies that illustrate cherubs bearing trophies of the gods. The Cupid and Psyche myth is based on the first novel that comes down to us from antiquity, known as the Metamorphoses (or The Golden Ass) of Apuleius, written in the second century but translated to Latin in 1469. The story, which was to become immensely popular, concerns the marriage of Cupid (mischievous God of Love) to the beautiful mortal Psyche (\"soul\") leading, after tribulations and trials, to the divinity of Psyche. The underlying theme of the story is that \"love conquers all.\" These Raphael frescoes illustrating the heavenly adventures of Cupid and Psyche are stunning, but for botanists and horticulturists an even greater treasure is found on the ribs enclosing the loggia's spandrels and severies. Here, the elaborate festoons and wreaths painted by Giovanni Martini da Udine (1487 22 Arnoldia 70\/2 The Villa Farnesina 23 coltura 2002). The resemblance of the maize images painted in Italy between 1515 and 1519 to races of maize in Spain, Portugal, and Italy is confirmatory evidence for the early origin of some of these races. There is direct evidence that maize seeds reached Rome in 1594. A letter from Peter Martyr D'Anghiera, an Italian teacher connected with the Spanish court, to Cardinal Ascoanio Sforza, vice-chancellor of the papal court, describes news of the early returning ships from the second voyages of Columbus and encloses seed of maize (McNutt 1912; Janick and Caneva 2005). Pome Fruits Five types of pome fruits are illustrated in the festoons: apple (Malus: 97 fruits of cultivated apple and 21 fruits of wild apple); pear (Pyrus: 78 fruits of European cultivated pear and 21 of wild pear); quince (Cydonia: 31 fruits), medlar (Mespilus: 27 fruits), and hawthorn (Crataegus: 30 fruits in two clusters). The number of images are indicative of the relative popularity of these fruits in Renaissance Italy. Of the apple fruits (Malus 24 Arnoldia 70\/2 The Villa Farnesina 25 Old World cucurbits seen in the festoons include watermelon (upper left) and various melons within Cucumis melo horticultural groups. Depictions of Lagenaria siceraria gourds include both the bottle type (with a broad, round base) and serpentine type (long, slender form). art, probably because of their phallic shape and association with the worship of Priapus, god of orchards and vineyards and the personification of the male generative organ (Morel 1984; Janick 2004). Finally, there are 3 images (9 fruits) of bitter gourds, known also as balsam apple (M. balsaminia) and balsam pear (M. charantis). Fruits are reddish and slightly warty, with a pointed end. The cucumbers in the paintings look like typical modern pickling types. There is also one image of squirting cucumber (upper left). Cucurbits (New World Species) Of particular interest for the festoon images are representatives of two species of New World cucurbits: Cucurbita maxima (fall and winter squashes and pumpkins) and Cucurbita pepo The cucumber images (13 groups, 25 fruit) all resemble the type known as \"American Pickling.\" There is a single image of the so-called squirting cucumber. Two types of Lagenaria fruit associated with white flowers are included: the inedible bottle gourd (var. fiasco) which is used largely for utensils, and the serpentine or club-shaped gourd (var. longissima), called cocuzza in Italy, that is edible when immature and still consumed in Sicily. There are 9 fruits of bottle gourds with slight variation in color and neck morphology. There are 19 groups (22 fruits) of cocuzza with subtle differences in shape based on the thickness of the calyx end suggesting that some might be hybrids of fiasco and longissima types. Cocuzza are widely displayed in Renaissance New World cucurbits seen in the paintings include large pumpkin or squash types of Cucumis maxima (upper left) and small gourd type examples of C. pepo. 26 Arnoldia 70\/2 The Villa Farnesina 27 lus is found in an illustrated manuscript, Livre d'Heures d'Anne de Bretagne, painted between 1503 and 1508 by Jean Bourdichon (ca. 1457"},{"has_event_date":0,"type":"arnoldia","title":"Maclura pomifera: Neither Apple Nor Orange","article_sequence":4,"start_page":28,"end_page":29,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25541","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d2708126.jpg","volume":70,"issue_number":2,"year":2012,"series":null,"season":null,"authors":"Hetman, Jon","article_content":"Maclura pomifera: neither Apple nor orange Jon Hetman T hough I confess that plants held little sway among my childhood interests, a few specific trees stand out in my early recollections, all due to their memorable fruits. A seemingly ancient apple tree in a neighbor's front yard was a climbing favorite, and bore small, mottled green fruits that were a delight to eat as long as you didn't overindulge. Behind my grandparent's garage in central Florida, a moss-covered orange tree provided fragrant spring flowers and slightly sour fruits that remain indelible sensations of my youth. But the fruits that perhaps fascinated me most belonged to an Osage orange tree that grew near my elementary school--large, hard as baseballs, and looking to us like green brains, the bumpy orbs with their citrusy aroma were a delightful mystery that inspired a number of ingenious games of our own design. In earlier days, Osage orange (Maclura pomifera) garnered significant interest among people of its native Oklahoma, Arkansas, Missouri, and Texas, though for very different reasons. Hunters in the Osage tribe fashioned war clubs and bows from the tree's bright yellow heartwood, which proved stronger than oak and as tough as hickory. Early settlers in the American frontier called it the hedge apple, planting it in thicket-like rows so that the thorny, interlacing branches sheltered fields from wind and provided an impenetrable animal barrier. As historian Paul Landacre famously described it, an Osage orange hedge was \"horse-high, bullstrong, and pig-tight.\" For me though, even as an adult, it's the fruits of these dioecious trees that really excite the imagination. If you cut one in half--and you may need a saw to do so--you'll discover a tough, pithy core surrounded by a couple hundred small seeds. Like other members of the Moraceae (mulberry family), Maclura bears a true multiple fruit composed of numerous separate ovaries, each developing from a separate female flower. In fact, the fruit's distinctive bumps--and their accompanying black, hairlike styles--rise from the fruit's numerous, tightly-packed ovaries. Though squirrels rip into fallen fruits to consume the seeds, noth- ing else seems to find these forbidding fruits the least bit appetizing. Nothing, that is, that still exists. In her book The Ghosts of Evolution, author Connie Barlow suggests that mammoths, mastodons, and other large herbivores of the North American plains ate Maclura fruits and were its dispersal agents before humans evolved their own interests in the plant. Although the exact details of the original collection of this monotypic species remain murky, we know that Maclura was among the botanical specimens gathered by Lewis and Clark on their transcontinental expedition of the American West. By studying saplings subsequently cultivated in the Philadelphia garden of Bernard McMahon, Constantine Samuel Rafinesque produced the first botanical description of the tree in 1817, naming it Ioxylon pomiferum, or \"poison apple.\" Perhaps unaware of Rafinesque's classification, Thomas Nuttall offered his own description the following year, honoring American geologist William Maclure with its generic epithet and proposing aurantiaca (\"orange colored\") as its specific epithet. Nearly a century later, Germany's Camillo Karl Schneider argued for the name that has stuck to this day, pairing Nutall's generic Maclura with a derivation of Rafinesque's specific, pomifera. Coincidently, this Teutonic connection to Maclura is compounded in the Arboretum's most spectacular accession (471-36-B) of the plant, a female obtained in 1936 from the Hermann A. Hesse Nursery of Weener, Germany. Growing today on a steep bank near the Centre Street wall across from Faulkner Hospital, the tree exhibits the criss-crossing, nearly horizontal branching that once made the species so desirable as a hedging plant. It is 36 feet (11 meters) tall and its two trunks have diameters at breast height of 14 and 16 inches (36 and 40.5 centimeters). Visit it in autumn, when its limbs bend beneath the weight of its fruits and its glossy leaves turn yellow, and you'll likely acquire fond associations of your own with this most singular of American fruiting trees. Jon Hetman is the Arnold Arboretum's Communications and Stewardship Officer. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23430","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd1608928.jpg","title":"2012-70-2","volume":70,"issue_number":2,"year":2012,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Constantine Rafinesque, A Flawed Genius","article_sequence":1,"start_page":2,"end_page":10,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25537","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270b36b.jpg","volume":70,"issue_number":1,"year":2012,"series":null,"season":null,"authors":"Mosquin, Daniel","article_content":"Constantine Rafinesque, A Flawed Genius Daniel Mosquin All pHotoS by tHe AUtHoR UnleSS otHeRWISe IndICAted V Stenanthium was first proposed as a subgenus of Veratrum L. by Asa Gray in 1837. Rafinesque had already suggested this group be recognized as its own genus, named by him as Anepsa, in 1832. When it was generally agreed upon that this group of species should indeed be considered its own genus, Rafinesque's earlier contribution was disregarded, and Gray's Stenanthium was conserved instead. Pictured here is Stenanthium occidentale A. Gray. iburnum rafinesqueanum--to a teenaged boy in Manitoba beginning to learn the scientific names of plants, this moniker stood out. Poa pratensis? Meadow grass or Kentucky bluegrass (pratensis = \"of a meadow\"). Caltha palustris? Marsh marigold (palustris = \"of a marsh\"). Aquilegia canadensis? Canada columbine or red columbine. Viburnum rafinesqueanum? Here was a mess of near-impenetrable letters, a poetic delight to my ears when recited, which I soon learned honored a man named Rafinesque. A few years later in a floristics lecture, the good-natured eye-rolling reaction of the professor to my question about Rafinesque started a broader curiosity about the man. Constantine Samuel Rafinesque was among the great American naturalists of the nineteenth century. He was also among the most controversial and eccentric natural history personalities of his time. In the course of four decades, he offended nearly every establishment botanist in the United States, leading to a disdain that persisted among these botanists and succeeding generations of their students. As one result, his contributions to botany and other natural history sciences were downplayed or ignored for many decades beyond his death in 1840. His reputation has been mended somewhat since the mid-nineteenth century, as those he interacted directly with passed away and several twentieth-century historians critically examined his life and work. What emerges is that the man was a flawed genius, whose inability to work within the bounds of scientific convention necessarily led to lower recognition than he would otherwise have deserved. Constantine Rafinesque 3 lIbRARy oF tHe ARnold ARboRetUM to immediately give an idea of Rafinesque and aspects of his personality, it is perhaps best to learn of his many roles in his own words: \"Versatility of talents and of professions, is not uncommon in America; but those which I have exhibited in these few pages, may appear to exceed belief; it is a positive fact that in knowledge, I have been a botanist, naturalist, Geologist, Geographer, Historian, poet, philosopher, philologist, economist, philanthropist ... by profession, a traveller, Merchant, Manufacturer, Collector, Improver, professor, teacher, Surveyor, draftsman, Architect, engineer, pulmist [one who treats pulmonary diseases], Author, editor, bookseller, library, Secretary ... and I hardly know myself what I may not become as yet: since whenever I apply myself to any thing, which I like, I never fail to succeed if depending on me alone, unless impeded and prevent by lack of means, or the hostility of the foes of mankind.\" RAFinesque's LiFe Rafinesque was turkish-born to a French father and a mother of German descent on october 22, 1783. He was reared in Marseilles, France, by his mother and his father's family; his father was a merchant trader who spent much time abroad. In 1792, his family fled to Italy to escape the French Revolution. A year later, his father died during a yellow fever epidemic in philadelphia. Rafinesque returned to France in 1797, where he remained until 1802. At the age of 19, he landed in philadelphia for three years, where his passion for botanizing the United States started immediately. He asserted that the brassicaceous Draba verna l. he picked up after stepping off the ship was a new species, as he generally believed that American counterparts of well-known european species could not be the same species. It is also in philadelphia where he began to write books and papers. In 1805, he returned to Italy where he resided for a decade (occasionally living under the name Constantine Samuel Rafinesque Schmaltz, in order to avoid anti-French sentiment). Here, he married in 1809, had a daughter born in 1811 and an infant son who perished in 1814. A return to the United States was made in 1815, though the boat he was traveling on was ship- Image of Constantine Rafinesque on the frontispiece of his 1815 publication Analyse de la nature. wrecked off long Island and he lost much of his collections and notes. Rafinesque lived in new york for three years, and helped to found the lyceum of natural History of new york. In 1818, a brief residence of under two years was made in philadelphia, before undertaking a posting as professor of natural History at transylvania University in lexington, Kentucky, from 1819 to 1826. post-professorship, he returned to philadelphia for the remainder of his life. on September 18, 1840, he died of stomach cancer. TAxonomiC ConTRoveRsies botanist, taxonomic scholar, and former director of the Arnold Arboretum elmer drew Merrill completed the voluminous Index Rafinesquianus in 1949 wherein he attempted the Herculean task of compiling the botanical work 4 Arnoldia 70\/1 Constantine Rafinesque 5 The genus Lomatium was proposed in 1819. It did not see much use for the next century, as critics declared it too close to the name Lomatia R. Br. (Proteaceae). Coulter and Rose briefly adopted it in a 1900 monograph of the Umbelliferae, but it didn't see widespread use until 1920 when Macbride pointed out that, according to the rules of botanical nomenclature, it was a valid generic name despite the similarity to Lomatia. Seen here, Lomatium brandegeei (Coult. & Rose) J.F. Macbr. is native to British Columbia and Washington. due to fire or other disasters), a nod to the principle of reproducibility in the scientific method. An additional, and critical, concept to understanding the controversy surrounding Rafinesque is that the linnaean system makes no attempt to define the boundaries of taxa. though a hierarchical framework is provided, the questions of \"What is a species?\" or \"What constitutes a genus?\" are left to the determination of taxonomists. this leeway gives the taxonomist much latitude in determining what might constitute a taxon. If the taxonomist errs on making too broad of a definition (i.e., \"lumps\" too much variability within a taxon), it increases the likelihood that her or his work will be revised by the next taxonomist to examine the taxon. Similarly, if the taxonomist errs on making too narrow of a definition (i.e., \"splits\" a group into separate taxa based on too little variability), the likelihood of revi- Parnassia glauca was one of a number of species published posthumously in 1840, in Rafinesque's Autikon Botanikon. 6 Arnoldia 70\/1 Constantine Rafinesque 7 reform, and called us Genera-mongers. We may in return call them Genera-shufflers, who want to squeeze plants into improper genera, and delay improvements by opposing the corrections of botanical blunders. It is to them that we owe the superfluity of synonyms: they often shuffle plants into 3 or 4 Genera, as linnaeus did for Heliopsis, until it must at last form a Genus of itself. It is a fact that almost all plants of doubtful Genera, are types of peculiar ones; the chances of it increase, as they are shifted.\" With the establishment of the linnaean system and the publication of Species Plantarum as the nomenclatural benchmark, linnaeus is credited with the valid publication of a large number of genera and species. linnaeus described about 1,440 genera, and most of these names are still in use today. by contrast, the splitter Rafinesque described approximately 2,700 genera--of these, no more than 50 or 60 are applied to recognized genera today (yet, had priority been applied, he would be credited with at least 160). linnaeus also generated almost 9,000 binomials (species names), and again, the large majority of these are in use today. Rafinesque did not quite match linnaeus in this category. of the 6,700 or so species names published by Rafinesque, fewer than 300 are generally accepted. Rafinesque's proclivity to deem the most minor variations as new species (and sometimes new genera) created work--much more work--for anyone who later attempted to publish a new species, write a monograph, or clarify names in a taxon. to give an example, Clintonia is a genus named by Rafinesque (and still recognized today). before Rafinesque erected a new genus for this group in 1832, its species were variously recognized as being in Dracaena (the first published name was in 1789), Convallaria, and Smilacina. According to The Plant List (drawing on information from the World Checklist of Selected plant Families), 41 names have been published within Clintonia (the actual number is likely higher). Working with a dataset of 35 names of \"High Confidence level\" (\"applied to the status of name records derived from taxonomic datasets which treat the whole of the taxonomic group in question on a global basis and have been peer reviewed\"), 30 are at the species rank (5 below the species level). Five of the species names are confidently recognized as \"Accepted\" species, and a single name for a recently described (1993) Asian species remains unresolved. the remaining 24 names are listed as synonyms, i.e., names that are considered to be already represented within the concept of a different name. of these 24 synonyms, 19 were published by Rafinesque. examples of species recognized by Rafinesque but generally regarded as minor variations within Clintonia uniflora (Sol.) Raf. include Clintonia angustifolia Raf. (a narrow-leaved entity), Clintonia biflora Raf. (a two-flowered entity), and Clintonia ciliata Raf. (presumably with fine hairs along the margins of an organ like a leaf or petal). If a taxonomist were to discover what she\/ he believes to be a new species of Clintonia, the taxonomic work involved would require at a minimum comparing it against the type specimens of other members of the genus and reviewing the taxonomic literature to ensure a previously published name and description (including all synonyms) does not conform to the purported new species. In practice, the taxonomist would further compare it against additional specimens of each species in order to properly account for variation within each species. In order to name a new species in Clintonia, the work required would involve reviewing all of Rafinesque's names and descriptions to determine if he had named the entity first. For a relatively simple group of species like Clintonia (5 accepted species), the task would be difficult in modern times, and very difficult at the time of Rafinesque. For more taxonomically complicated genera, like Trillium, Rafinesque made the difficult near-impossible. there are about 38 recognized species of Trillium in north America, with more than two-thirds of these from eastern north America. Rafinesque is presently responsible for 3 of these accepted names, though he described an additional 31 species and 67 varieties. this onslaught of published names of additional genera and species in many eastern north American plant groups, sometimes poorly described, was not well received. Amos eaton, a botanist and author of the 1817 Manual of Botany for the Northern States, was generally 8 Arnoldia 70\/1 Constantine Rafinesque 9 sympathetic to Rafinesque and considered him a friend. However, in 1817, he wrote to his student John torrey: \"I am glad Mr. Rafinesque has not set you all wild. Why can not he give up that foolish european foolery, which leads him to treat Americans like half-taught school boys? He may be assured, he will never succeed in this way. His new names with which he is overwhelming the science will meet with universal contempt.\" eaton accurately predicted the ultimate approach by much of the botanical establishment--ignore much of Rafinesque's work, to the extent that the principle of priority was overridden in many cases to exclude Rafinesque's contributions. Asa Gray, the pre-eminent American botanist of the nineteenth century, contributed to the practice of discounting Rafinesque. though he was charitable towards Rafinesque's earlier work, Gray's influence cemented the rejection of Rafinesque's ideas about new genera and species when he wrote the following about Rafinesque after his death: \"Many of Rafinesque's names should have been adopted; some as a matter of courtesy, and others in accordance with the strict rule.... one who, like Rafinesque, followed the easy rule of founding new genera upon all these species, could not fail to make now and then an excellent hit; but as he very seldom knew the plants themselves, he was unable to characterize his proposed genera, or to advance our knowledge respecting them in the slightest degree. In his later publications, this practice is carried to so absurd an extent as entirely to defeat its object ... A gradual deterioration will be observed in Rafinesque's botanical writings from 1819 to about 1830, when the passion for establishing new genera and species, appears to have become a complete monomania\". Originally published as Saxifraga ranunculifolia by Hooker in 1832, Rafinesque clearly disagreed. He erected the genus Hemieva in 1836 to segregate this species, but was ignored. In 1879, Asa Gray named a new genus and species Suksdorfia violacea. In 1891, H. Engler assigned Saxifraga ranunculifolia to Suksdorfia. Recent phylogenetic studies suggest Suksdorfia ranunculifolia is within a distinct genus, so Hemieva ranunculifolia (Hook.) Raf. was resurrected in the second edition of The Jepson Manual (2012), a major taxonomic reference. in the lapse of time,\" and that \"every variety is a deviation which becomes a species as soon as it is permanent by reproduction.\" Rafinesque's ideas were informed by Adanson from 1763, to whom he gives credit: \"Adanson ... was like linnaeus, necker and myself (in fact like all acute observers) a strenuous supporter of the doctrine that Species were unlimited, and increasing by the natural process of semination, deviation, variation, hybridization and such. Whence he concluded that we could hardly ascertain the primitive types of species, that many known to ancient botanists were lost or no longer found, while new ones were evolved in mountains, groves, fields, and gardens.\" on evoLuTion Another area where Rafinesque generated controversy was in his ideas about how species and genera were formed. one of the reasons Rafinesque named so many species and genera was because (in his own words, from 1832), \"the truth is that Species and perhaps Genera also, are forming in organized beings by gradual deviations of shapes, forms and organs, taking place 10 Arnoldia 70\/1 "},{"has_event_date":0,"type":"arnoldia","title":"A Quarter-Century Perspective on the Center for Plant Conservation Collections at the Arnold Arboretum","article_sequence":2,"start_page":11,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25535","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270af6d.jpg","volume":70,"issue_number":1,"year":2012,"series":null,"season":null,"authors":"Hird, Abby","article_content":"A Quarter-Century Perspective on the Center for Plant Conservation Collections at the Arnold Arboretum Abby Hird Flowers and fall foliage of Amelanchier nantucketensis. NANCy RoSe G iven the array of current threats to biodiversity (including habitat destruction, pollution, climate change, and invasive species), it is no surprise that roughly one out of every three plant species in the world is threatened with extinction. As native habitats are changing and disappearing, ex situ conservation (preservation of species outside their natural habitat as living plants, seeds, or other viable tissue) efforts are even more vital to the successful conservation of plants. Public gardens, numbering more than 700 in the United States alone (BGCI 2012), offer valuable resources, facilities, and horticultural expertise that support conservation efforts. These ex situ refuges also allow visitors a unique chance to learn about and observe threatened species firsthand. While these important plant collections serve as insurance policies against extinction for many species, the recent North American Collections Assessment found that only 39% of North America's threatened species are currently cultivated in public gardens; clearly, there is much opportunity to increase rare plant conservation collections (Kramer et al. 2011). In addition to increasing the number of threatened species in ex situ collections, broadening genetic diversity within those collections will support meaningful conservation applications. During my Putnam Fellowship from 2008 to 2010 I worked with Arboretum Curator of Living Collections Michael Dosmann to assess the conservation potential of the The ARNoLD ARBoReTUM 12 Arnoldia 70\/1 MIChAeL DoSMANN Center for Plant Conservation 13 CPC ColleCtions MAnAgeMent guidelines (CPC 2007) 1. Taxa should be proposed and accepted by the CPC Science Advisory Council for inclusion into the National Collection. 2. Propagative materials should be collected from the wild in accordance with CPC guidelines and should be maintained in protective storage. 3. A usable seed storage and germination protocol should be developed for the taxon and initial seed viability should be determined if possible. 4. horticultural techniques for ex situ cultivation should be established and documented, and the taxon should be successfully raised to reproductive maturity. 5. Adequate propagules and data should be stored in at least two separate secure sites 6. An initial baseline germination test should be conducted on stored seed accessions of the taxon, and viability should be retested at appropriate intervals, using enough seed if possible to detect statistically valid declines in viability. 7. Collaborative research agreements to be established for taxa as necessary and appropriate. 8. Legitimate reintroduction programs or experimental reintroductions are encouraged. In total, the Arboretum has assisted in the conservation efforts of 24 threatened CPC species by collecting wild germplasm and maintaining those plants in the living collections MIChAeL DoSMANN Fothergilla major bears fragrant, bottlebrush-like flowers in spring. 14 Arnoldia 70\/1 Center for Plant Conservation 15 About natureserve g-ranks and threat levels NaTUReSeRve'S Global Conservation Status Ranks (G-ranks) are the most comprehensive source of conservation information on species native to the United States or Canada (NatureServe 2012). G-ranks can be used to gauge the \"level of need\" for each species, which is useful when prioritizing collections curation and development activities such as repropagations, voucher collection, or backup germplasm distribution at an institution. Thus, Torreya taxifolia with a G-rank of G1 (Critically imperiled) has the greatest conservation need (the most threatened in the wild, with the fewest remaining wild populations) among current CpC species and first priority in collections management decisions; while Diervilla sessilifolia with a G-rank of G4 (apparently Secure) has a relatively lower conservation need. Global Rank Categories GX: presumed extinct, GH: possibly extinct, G1: Critically imperiled (5 or fewer populations remain), G2: imperiled (very few remaining populations), G3: vulnerable (relatively few remaining populations), G4: apparently Secure (common; widespread and abundant), Q: Questionable Taxonomy, GNR: Unranked, GNa: Not applicable (see Table 1). A majority of the Arboretum's CPC collections began via collecting expeditions to the southeast and northeast regions of the United States in the late 1980s and early 1990s by Rob Nicholson, then plant propagator for the Arnold Arboretum (Nicholson 1996). At the time of this assessment there were 13 species assigned to the Arnold Arboretum. Many of the current and historic CPC species collected by the Arboretum originate from the southeastern United States, and some have experienced cold hardiness issues in the Northeast. In the past 15 years, due to significant decline in health and numbers of living accessions, about half of the original CPC species have been transferred to more appropriate institutions closer to their native range and with more compatible climates. balsam wooly adelgid (Adelges piceae) introduced from europe. In 1876, Asa Gray first collected a wild plant for the Arboretum (accession 1522), which did not survive. Since then, several specimens were unsuccessfully introduced to the the Arboretum. A collecting trip in 1985 supported the establishment of the CPC collection of this species, which have suffered excessive losses due to incompatible climate and spider mite infestations. The Arboretum maintains 6 specimens from 3 states (Virginia, North Carolina, and Tennessee). Amelanchier nantucketensis: The Nantucket shadbush is a stoloniferous shrub which forms dense colonies in its restricted native habitat along the northeastern Atlantic coast. Flowers usually open in May but are small and hard to notice. Threats to this species include overcrowding by other plant species, harmful management practices such as fire suppression, and uncontrolled land development of coastal habitat. The Arboretum maintains specimens collected in the 1980s from New york, Massachusetts, and Maine. A large group was successfully transplanted during the Bradley Rosaceous Collection renovations in 2009 and now thrives near Dawson Pond. PlAnts in tHe Arnold ArboretuM's CPC ColleCtions Abies fraseri: The Fraser fir is well-known in the Christmas tree industry due to its spirelike crown and fragrant foliage. Reaching heights of up to 25 meters (82 feet), this species is native to the Smoky Mountain Range and is unique because it grows at high elevations. It is severely threatened in the wild by the invasive 16 Arnoldia 70\/1 Center for Plant Conservation 17 Rhododendron prunifolium: The plumleaf azalea is one of the showiest native azaleas, and may reach up to 6 meters (19.7 feet) tall in the wild. It has glabrous leaves and bears clusters of red-orange flowers in July and August. It is native to Alabama and Georgia and is threatened by logging and low seedling numbers in the wild. The Arboretum currently has specimens from two locations in Georgia, and is responsible for introducing this species into cultivation in the early 1900s via plant collector T. G. harbison. Rhododendron vaseyi: The pinkshell azalea is an upright shrub known to grow up to 5 meters (16.4 feet) tall in the wild. Scentless (and frost resistant) pink flowers emerge in April prior to leaf bud break, providing striking ornamental value. This species is native to North Carolina, and is threatened by land development and illegal collecting in the wild. The Arboretum introduced it to cultivation in 1880, and maintains several specimens from North Carolina which thrive in the Boston climate. Spiraea virginiana: The Virginia meadowsweet is a 1- to 2-meter-tall (3.3 to 6.6 feet) shrub that forms dense clumps of upright, arching stems with cream colored inflorescences in May. This species is endemic to the central and southern Appalachians, where its sporadic populations are threatened by competition with fast growing herbs and vines, habitat destruction including dam construction, and lack of sexual reproduction. Plants were first collected by the Arnold Arboretum in 1919 by T. G. harbison in a Fraser fir cone. North Carolina (accession 10160), grown at the Case estates, and then repropagated via cuttings and brought to the main Arboretum grounds in 1988, where the lineage still exists today. When this CPC collection was established in the mid1980s and 1990s, the Arboretum amassed one of the most extensive ex situ collections of this species in the world, composed of plants from all states where it is currently known to grow. Two groups were recently transplanted to beds near the South Street and Mendum Street gates. Torreya taxifolia: once a towering tree of 15 meters (49.2 feet) or more, the stinking cedar (named for its pungent, sharp needles) is native to Georgia and Florida, and is now one of the most threatened conifers in the world because of a fungal disease. The few remaining wild individuals have been reduced to root suckers.Until 2010, the Arboretum maintained 33 specimens from known remaining populations. BILL CooK, MIChIGAN STATe UNIVeRSITy, BUGWooD.oRG Magnolia pyramidata: The pyramid magnolia grows 3 to 7 meters (9.8 to 23.0 feet) tall, and produces creamy white flowers that give it potential as an ornamental landscape plant. It is native to a limited range along the coastal plain of the southern and southeastern United States, and is threatened by land development. The Arboretum does not currently have specimens in the living collections, but had grown two lineages from Texas that were removed in 2001 when it was determined that the specimens were not M. pyramidata. 18 Arnoldia 70\/1 Center for Plant Conservation 19 CoURTeSy oF TRoy UNIVeRSITy heRBARIUM, ALABAMA PLANT ATLAS WeBSITe (hTTP:\/\/FLoRAoFALABAMA.oRG) Magnolia pyramidata has a limited native range in the United States. 20 Arnoldia 70\/1 NANCy RoSe Center for Plant Conservation 21 table 2. recommended number of Populations in a rare Plant sampling Program for Capturing genetic diversity at the Population level (Falk and Holsinger 1991) Number of extant populations 1 2 3 4 5 >5 Number of populations Sampled 1 2 3 3-4 3-5 4-5 tee 2007). When comparing the total number of living accessions to the total number of living lineages, each CPC species is represented by 1 to 2 accessions per lineage. This assessment showed that both of the Diervilla species had a higher number of accessions per lineage, demonstrating redundant clones within the same lineages. To maintain appropriate accession-to-lineage ratios for the CPC collections, we identified lineages and accessions that could be bulked up via clonal propagation and others that could be \"thinned\" by sending back-up material to other institutions. Rhododendron prunifolium bears red-orange flowers with prominent red stamens. Plants health conditions through time and total numbers of living plant specimens give an indication of how well a species grows in the Arboretum and can provide guidance for collections management. At the time of this assessment in 2009, most CPC specimens were healthy. however, management needs were further considered for species with significant proportions of specimens in fair or poor condition, such as Torreya taxifolia, Amelanchier nantucketensis, and Abies fraseri. Also, Magnolia pyramidata, with no living plants represented in the collection, was prioritized for a collection transfer or germplasm acquisition. By using the Arboretum collections standard of maintaining an average of 2 plants per unique accession (Living Collections Committee 2007), we identified collections redundancy or deficiency for each CPC species. As we analyzed accession-to-lineage ratios, we also compared the total number of living accessions with the total number of living plants CoURTeSy oF MT. CUBA CeNTeR Finding a Home for Torreya taxifolia ToRReyA TAxIfoLIA, once a towering giant in the forests of Georgia and Florida, has been diminished to twig-like sprouts by an obscure fungal disease over the past century. It is now one of the most threatened conifers in the world. Several ongoing conservation efforts strive to understand the pathology of the disease and find effective management and reintroduction strategies. Several ex situ collections of the species have been aimed at conserving the narrowing genetic diversity of extant wild populations as well as producing seeds and cuttings for research. A large-scale ex situ effort began in 1985, funded by the CPC and the Arnold Arboretum (Nicholson 1996). Rob Nicholson and Mark Schwartz collected cuttings from 163 wild lineages of T. taxifolia and then distributed resulting plants to 10 institutions in North America and europe in the early 1990s. Using a 1996 Arboretum inventory of 156 of the original lineages as a foundation, we conducted an international inventory of this species in 2009 and tracked down all possible specimens that originated from the original CPC material. Fortunately most of the lineages had been preserved among the institutions surveyed (a benefit of backing up collections). But about 20% were represented by only one or a few remaining plants per lineage, and about 40% of lineages existed only at one or two institutions. Lessons learned from this long-term ex situ effort include ensuring a collection holder has appropriate horticultural know-how, climatic compatibility, and staff commitment for successfully maintaining a collection. For example, a loss of 70% of unique lineages at the Arnold Arboretum was observed from 1989 to 2009. This loss is attributed to incompatible climate, poor adaptability to container nursery conditions, and human error (staff changeover, labeling errors, etc.). Further, 5 of the 8 institutions still maintaining the original T. taxifolia germplasm required accession data cleanup and several specimen identifications were determined lost or unknown due to accidental dissociation with accession numbers, labels, or records. This long-term ex situ conservation effort dem- Foliage of Torreya taxifolia. onstrates how living collections can contribute to the collective conservation power of public gardens. As a result of the 2009 ex situ inventory for T. taxifolia, redistribution of germplasm has occurred among collection holders to preserve and back up ex situ maternal lines at multiple institutions. Further, this inventory led to a successful transfer of this important CPC collection from the Arnold Arboretum to the Atlanta Botanical Garden in 2010. ReBeKAh D. WALLACe, UNIVeRSITy oF GeoRGIA, BUGWooD.oRG Center for Plant Conservation 23 per species, also taking into account specimen health. This allowed us to identify specific plants in need of repropagation, removal, or relocation. one particularly successful example of making management decisions to improve plant health is the Amelanchier nantucketensis specimens in the Bradley Rosaceous Collection (BRC). Poor health had been recorded for these plants for several years, and during bed renovations in the BRC they were transplanted to new beds near Dawson Pond. This location's higher soil moisture has resulted in improved health for the plants. A common issue identified for Amelanchier nantucketensis, both Diervilla species, and Spiraea virginiana was maintaining individuals of these mass-forming species. As a result, these specimens were put on a pruning schedule to prevent uncontrolled spreading and suckering. Since the Arboretum's primary goal with the CPC collections is preservation of living germplasm, long term survival of the CPC plants is a top priority. Collections management at the Arboretum includes the preservation of unique lineages through clonal repropagation if needed. Sometimes plants brought to the Arboretum are not well-suited to survive in the collections for reasons such as lack of compatibility to cultivation or the local climate. Species whose records show high levels of lineage or plant loss, such as Torreya taxifolia, likely represent poor compatibility with Arboretum conditions, making them potential candidates for transfers to institutions better able to cultivate them. Additional supporting documentation may include observations, voucher herbarium specimens, images, verifications, and recorded instances of collections use (for tours, publications, and educational projects involving a species). herbarium specimens and images offer long-term genetic and biological information that can enhance understanding and aid in conservation of a threatened species. The Arnold Arboretum Cultivated herbarium sets a goal to document the living collections with vegetative, flowering, and fruiting material per unique lineage (Curatorial Department 2009). This CPC assessment identified gaps in passport data and supporting documentation for each species. In addition to augmenting geographic passport data for many CPC accessions, we also established herbarium specimen and image collecting targets, as well as past verifications that could be entered into the plant records database. ColleCtions enHAnCeMent Priorities The Arboretum has taken a number of positive steps following this assessment to improve and more effectively manage the CPC collections, making them more valuable and accessible for research, education, and conservation. Individual species reviews allowed us to create a prioritized master list of recommended curatorial and horticultural actions based on collections goals and needs. Accomplishments include enhancements in plant records information through the addition of county names, latitude and longitude, or other location information when possible. Voucher and image collection has also been a priority for the curatorial department, and over 350 herbarium specimens have been collected to further document the CPC collections. Recommended repropagations, removals, and relocations have been completed, including repropagation of two Abies fraseri specimens which are failing in the collection; addition of new lineages (Rhododendron vaseyi); removal of non-wild-origin plants and acquisition of new wild-origin lineages (Rhododenron prunifolium); planting out of nursery stock (Torreya taxifolia); and removal of redundant specimens Supporting Documentation The geographic, temporal, and environmental details about the source of an accessioned living plant are referred to as the passport data, which are curated in the Arboretum's plant records. Passport data can make collections more valuable for conservation, education, horticulture, and research by associating valuable habitat or biological information with each specimen. For wild-collected plant material the value of a collection increases with the amount of passport data. This can range from coarse geographic information such as country and state to highly local information such as soil type or altitude of an original collection location. 24 Arnoldia 70\/1 MIChAeL DoSMANN Center for Plant Conservation 25 Spiraea virginiana sources revealed through molecular study iN THe SUmmeR of 2008, leaf tissue samples of all living specimens were sent to Jessica Brzyski, then phD candidate at the University of Cincinnati, who was researching reproduction of S. virginiana. in the summer of 2009, Jessica visited the arboretum as a Deland award recipient and conducted controlled pollinations to determine the level of self-compatibility and out-crossing ability for S. virginiana. She was able to provide a summary of her molecular studies using the leaf tissue, which provided clarification on some of the questionable paternities of a few living specimens at the arboretum, some of which had grown into large masses in recent years. The controlled pollinations were inconclusive, as 2009 was an extremely rainy summer and most of the pollinations were completed in the rain. However, the molecular information helped us re-identify specimens that came from the same populations as other specimens with known identities. USDA-NRCS PLANTS DATABASe species would bring conservation work closer to home and likely result in increased success for threatened species grown, maintained, and utilized at the Arnold Arboretum. References BGCI. 2012. GardenSearch database. Accessed online: www.bgci.org\/garden_search. php. Botanic Gardens Conservation International. CPC. 1984. The Center for Plant Conservation Archives. The Arnold Arboretum of harvard University. CPC. 2007. Management Guidelines for Participating Institutions. Participating Institution handbook. CPC. Curatorial Department. 2009. Cultivated herbarium Collection Policy. The Arnold Arboretum of harvard University. Falk, D. A. and K. e. holsinger (eds.) 1991. Genetics and conservation of rare plants. New york: oxford University Press. hird, A. and M. Dosmann. 2010. CPC Collections Analysis. The Arnold Arboretum of harvard University. horticulture Department. 2012. Landscape Management Plan, 2nd edition. The Arnold Arboretum of harvard University. Kramer, A., A. hird, K. Shaw, M. Dosmann, and R. Mims. 2011. Conserving North America's threatened plants: Progress report on Target 8 of the Global Strategy for Plant Conservation. Botanic Gardens Conservation International U.S. Living Collections Committee. 2007. Living Collections Policy. The Ar nold Arboretum of harvard University. NatureServe. 2012. NatureServe Conservation Status. Accessed online: http:\/\/www. natureserve.org\/explorer\/ranking. htm Nicholson, R. 1996. CPC file archives. The Arnold Arboretum of harvard University. Abby hird is a Research Associate with Botanic Gardens Conservation International's United States office, based at the Arnold Arboretum. Spiraea virginiana from Britton and Brown's An illustrated flora of the northern United States, Canada, and the British Possessions, 1913. "},{"has_event_date":0,"type":"arnoldia","title":"Book Review: Conifers Around the World","article_sequence":3,"start_page":26,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25536","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270b326.jpg","volume":70,"issue_number":1,"year":2012,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Book Review: Conifers Around the World Peter Del Tredici Conifers Around the World Zsolt Debreczy and Istv Book Review 27 tions of the book cover species ranges (474 maps reproduced from other sources); lengthy descriptions and numerous photographs of typical conifer habitats, arranged by continent; and a highly unusual \"bark gallery\" consisting of 648 color photos that augment the photos in the species treatments. And finally, there is a 130page introduction that describes the history, morphology, ecology, taxonomy, biogeography, and evolution of conifers along with a complete listing and description of conifer families (written by Robert Price). This introduction could easily be expanded into a stand-alone book about conifer morphology and natural history. An early version of Conifers Around the World was published in Hungarian in 2000 (Fenyk a F 28 Arnoldia 70\/1 Book Review 29 JEFF BISBEE This specimen of Cupressus arizonica var. arizonica displays two types of bark: scaly, persistent ribs on the main trunk, and detaching leathery plates on the lateral branches. spatial separation of closely related species (e.g., those within Taxodium) that are growing in separate regions. The taxonomy used in the book, for the most part, follows the accepted botanical classification used by most conifer authorities, but Debreczy and R "},{"has_event_date":0,"type":"arnoldia","title":"Dipelta floribunda: A Shrub of Subtle Beauty","article_sequence":4,"start_page":32,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25538","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270b76f.jpg","volume":70,"issue_number":1,"year":2012,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Dipelta floribunda: A Shrub of Subtle Beauty Michael Dosmann I n the horticultural world, it is not uncommon to hear plant lovers laud a particular plant's endless weeks of flowering, months of dazzling autumn leaf color, and flamboyant, persistent fruits the size of golf balls. I can appreciate plants like that, and yet sometimes I want something subtler. English novelist George Meredith wrote that \"Speech is the small change of silence,\" and as I apply that maxim to the garden I find that I am drawn toward plants that possess quiet interest. One such plant is Dipelta floribunda, the rosy dipelta, a shrub native to central and western China. The Arboretum has cultivated rosy dipelta for over a century, the first seeds coming to the Arboretum in February 1911 from E. H. Wilson's collection from Fang Hsien, western Hubei, the previous October. He made the collection from plants growing in \"sunny places\" at altitudes of 1,200 to 1,800 meters (3,937 to 5,905 feet). Seventy years later, the Arboretum received its latest accessions of this species, collected in Hubei during the 1980 Sino-American Botanical Expedition. Plants from two separate accessions from the 1980 SABE grow in the Arboretum, as does one large plant (accession 14514-B) from the Wilson accession. The Explorers Garden atop Bussey Hill serves as perhaps the best place to see these plants, though there is also another fine mass planting of rosy dipelta along Peters Hill Road. Dipelta floribunda bears fragrant, pinkishwhite flowers, typically blooming in early May. Each tubular corolla comprises five fused petals, with the two upper lobes forming a top lip, while the basal three lobes form a lower lip. Yellow pigment splashes along the lips and throat of the flower, no doubt serving as nectary guides for the bees that pollinate the flowers. At the base of the corolla are greenish bracts that increase in size as the growing season advances, surrounding the fruits (two-seeded achenes) as they mature. The round, 1-inchwide, papery bracts provide a bit of late summer interest--particularly as they blush a tawny pink--and also aid in the wind dispersal of the seeds. A casual examination of Dipelta reveals similarities with Kolkwitzia amabilis, beauty bush, also introduced by Wilson. The two genera are closely related to each other within Caprifoliaceae, the honeysuckle family; their flowers look similar, though Kolkwitzia fruits have but a single seed and lack Dipelta's papery bracts. Rosy dipelta is a large, vase-shaped shrub that typically attains a height of 12 to 15 feet (3.7 to 4.6 meters) and a width of 6 to 8 feet (1.8 to 2.4 meters). The leaves are lanceolate and rather coarse, and tend to abscise in the autumn with little effective color change. But when they do drop from the plant they reveal another bit of quiet interest. With a few years of age, the bark of the stems begins to shed in long, vertical, tawny-white strips. In the garden, some may think this somewhat messy (plantsman Michael Dirr muses that \"the entire matrix ... assumes the presence of a pile of sticks\"). However, I like this trait for both its tactile quality and its visual appeal in the winter. An undulating row of several of these fine shrubs at the back of a mixed perennial border provides an excellent backdrop, particularly when they are pruned to remove lower branches. Mix in several beauty bushes to extend the flowering season a few weeks, add a Heptacodium miconioides (seven-son flower) to provide late summer blooms, and enjoy all three of them for their habit and bark interest. Michael Dosmann is Curator of Living Collections at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23429","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160856e.jpg","title":"2012-70-1","volume":70,"issue_number":1,"year":2012,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Japanese Flowering Cherries A 100-Year-Long Love Affair","article_sequence":1,"start_page":2,"end_page":14,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25533","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270ab6f.jpg","volume":69,"issue_number":4,"year":2012,"series":null,"season":null,"authors":"Aiello, Anthony S.","article_content":"Japanese Flowering Cherries --A 100-Year-Long Love Affair Anthony S. Aiello T his year marks the 100th anniversary of the 1912 planting of the famous flowering cherries surrounding the Tidal Basin in Washington, D.C. The story of how they came to be planted is worth exploring, given the centennial anniversary, the lasting impact of the planting efforts, and the continued public fascination with flowering cherries. Although the Tidal Basin plantings seem like a singular event, the interest in flowering cherries was widespread in the early 1900s, and these plants came into the United States through a number of different sources. Around this time both the USDA's Office of Foreign Seed and Plant Intro- duction (under David Fairchild) and the Arnold Arboretum were instrumental in bringing many cultivated varieties into the United States as part of a broad interest in flowering cherries. Based largely on the efforts of Fairchild, Charles S. Sargent, and E. H. Wilson, there was a surge in the number of varieties available in the first quarter of the twentieth century. The flowering cherries, or sakura, have been an integral part of Japanese culture for centuries. \"Japanese flowering cherries\" is a general term for a taxonomically complex group of plants that includes several well-known taxa such as Prunus subhirtella (Higan cherry), PETEr DEl TrEDICI The famous flowering cherry trees around the Tidal Basin in Washington, D.C. Japanese Flowering Cherries 3 ANTHONY S. AIEllO Prunus 4 Arnoldia 69\/4 lIBrArY OF CONGrESS PrINTS AND PHOTOGrAPHS DIvISION Japanese Flowering Cherries 5 MATTHEW JONES ArCHIvES OF THE ArNOlD ArBOrETUM with insects and diseases. All 2,000 trees were burned and, as can be imagined, this created a great deal of diplomatic consternation. Fortunately this was all overcome and a second shipment of 6,000 insect- and disease-free trees reached the United States in 1912. One half of these were sent to New York City, where some of the original Yoshino cherries grow near the reservoir in Central Park. The better known half of this shipment were the 3,020 trees that were sent to Washington and were planted Yoshino cherry blossoms frame the Jefferson Memorial in Washington, D.C. around the Tidal Basin, on the White House grounds, and in other areas in the city, where they quickly made the capital famous for its cherry blossom displays. These original trees were made up of 11 varieties of Prunus serrulata (1,220 plants) and 1,800 plants of Yoshino cherry (Prunus 6 Arnoldia 69\/4 Japanese Flowering Cherries 7 serrulata `Shirotae') cherries from the same to importing, growing, and hybridizing flowsource in 1912. ering cherries (Buchan 2011). His 1948 book, Flowering cherries continued to be very popuOrnamental Cherries, was responsible for lar between the World Wars. One of the leading spreading the gospel of growing cherries both proponents and sources of flowering cherries in the United Kingdom as well as on the Conwas Anton Emile Wohlert, the proprietor of tinent (Ingram 1948). If you happen to visit the the Garden Nurseries in suburban Philadelphia Philadelphia Flower Show or tour the city in (Wister 1955 Cherries in print AN INDICATION of the popularity of flowering cherries can be gained by reviewing the Arnold Arboretum's Bulletin of Popular Information and its successor, Arnoldia (Del Tredici 2011). Flowering cherries were mentioned as early as 1911, and their virtues were extolled regularly from the 19-teens through the 1930s (for examples, see Bulletin of Popular Information: New Series, vol. III (3) May 14, 1917: pp. 9 Japanese Flowering Cherries 9 newly added plants. In addition to trees dating tried on more of our mature cherry trees, with to the Morris Estate era, there were continual very similar results. waves of cherry varieties accessioned from the What began as trial-and-error attempts has 1940s through the 1980s. In the 1940s we received evolved into a regular retrenchment or restoraa large consignment of trees from the Scott Arbotion pruning program, based on the ideas estabretum, including a few that remain today. These lished in Europe for veteran tree management were followed by a group of plants from Kings(Fay 2002). We begin the process of targeted ville Nursery in the late 1950s, from Princeton pruning by reducing the end-weight of declining Nurseries in the mid-1960s, and more cultivars and decaying older branches. Major portions of from the U.S. National Arboretum in 1983. these branches are removed, lessening the endOne often reads that cherries are short-lived, load on these branches and reducing the risk surviving for not more than 50 or 60 years, so of failure along with hazards to the public and it may be surprising to learn that we have cherry trees that were planted by John and lydia Morris prior to the establishment of the Morris Arboretum in 1932. Our collection has individuals up to 100 years old because we use specific management practices for veteran trees. We work with the natural life cycles of these trees, managing them for longevity and safety and rethinking our approach to arboricultural practices. By implementing the practices of veteran tree care, we have been able to prolong the lives of our old flowering cherries almost indefinitely (Fay 2002). I could say that we began this process through careful literature research and a prescient understanding of veteran tree biology, but the reality is more serendipitous than that. In the early 1980s, then Morris Arboretum curator Paul Meyer (now our director) began to rejuvenate our Prunus collection by removing older trees and replanting with newly propagated plants that we had received from the National Arboretum. A 1940s accession of Prunus 10 Arnoldia 69\/4 ANTHONY S. AIEllO Japanese Flowering Cherries 11 ANTHONY S. AIEllO example of this is an old specimen of Prunus subhirtella `Pendula' that was planted prior to 1932. This tree has a highly decayed trunk with a band of healthy bark and one large remaining branch. For the past few years we have removed all but about five of these sprouts and are encouraging the basal rejuvenation of these to form a new tree (Fay 2002). Eventually we will remove all but one or two of these and then allow the original trunk to decay completely. Cherries have an especially interesting biology because of their tendency for endocaulous rooting, a process of forming roots from portions of stem tissue; these roots result in a successional trunk as they grow down through the decaying parent trunk (Fay 2002; liu and Wang 1992). As the inner trunks of older plants decay, often there is a shell of living tissue surrounding a core of rich decomposed organic matter from the old wood. The tree often initiates roots into this rich medium, and as root tissue grows down through the core of the tree, it provides added structural support to Management of the vigorous new sprouts around this old Prunus subthe tree's upper portions (Jenik 1994). hirtella `Pendula' allows the specimen to be rejuvenated. This process is especially apparent in old flowering cherries, and an extreme to stand on its own, but with about four feet example occurred with another of our old of above-ground root tissue forming the new Prunus subhirtella `Pendula' plants, in this trunk. Since then this tree has continued to case a plant that is shown on our 1909 Atlas prosper, a lazarus of a plant having returned of Compton (the Morris Estate). In the mid from the brink. It now grows vigorously across 1990s this tree was in significant decline, from our visitor center, providing a fabulous with a severely decayed old trunk supporting spring display. a few feeble branches. For a number of years Future eFForts with FLowering we observed a major root growing within a Cherries cavity in the trunk and leafy shoots arising A few years ago I began to expand the Morris from the top of this root with increasing vigor. Arboretum's cherry collection by propagating With each passing year the root became more early- and late-flowering varieties to extend the trunklike as the old trunk further deteriorated period of flowering interest. Before this project until it was a standing hollow shell. In the began I was intimidated by Prunus propagafall of 1997, the old rotting trunk simply fell tion, believing that, like many other rosaceous to the ground under its own weight. We were plants, they had to be grafted or budded to be delighted to see that the \"new tree\" that had reproduced. Fortunately our propagator, Shelformed inside of this shell was strong enough 12 Arnoldia 69\/4 Japanese Flowering Cherries 13 Arboretum. last year we rooted cuttings of these and look forward to growing and distributing them. `Gyoiko' is an especially interesting plant because it has chartreuse flowers with thin pink and white streaks in the center of the petals. The name translates as \"colored court-robes\" and refers to the green, white, and purple robes of women in the ancient Japanese imperial court (Kuitert 1999). In the original 1912 Washington planting, all 20 specimens of `Gyoiko' were planted at the White House. `Jo-nioi' (\"supreme scent\" or \"firstclass fragrance\") has single white flowers that bloom in profusion and is known as one of the most fragrant of the flowering cherries (Kuitert 1999). Although once more commonly grown, it has vanished from our landscapes and would make a fine addition to any garden. The flowering cherries at the Morris Arboretum are a prime example of how a living collection can fulfill multiple aspects of our mission, namely, collections preservation, horticultural display, research, and education. The cherry collection Three Prunus serrulata cultivars, including the unusual green-flowered `Gyoiko', are is a model for preserving our featured on this page from a 1916 14 Arnoldia 69\/4 "},{"has_event_date":0,"type":"arnoldia","title":"Charlie Deam and the Deam Oak (Quercus x deamii)","article_sequence":2,"start_page":15,"end_page":21,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25532","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270a76b.jpg","volume":69,"issue_number":4,"year":2012,"series":null,"season":null,"authors":"Hibben, George","article_content":"Charlie deam and the deam oak (Quercus x deamii) George Hibben SpuRgeOn-gReene phOTOgRAphS, ARChIveS AnD SpeCIAL COLLeCTIOnS, BALL STATe unIveRSITy LIBRARIeS A handy reference in the Arnold Arboretum's curatorial office is a paperback reprint of Trees of Indiana, originally published by the Indiana State Board of Forestry in 1912. It was written by Charles Clemon Deam (1865 16 Arnoldia 69\/4 Quercus x deamii 17 Accession 7033 e 7033 F 21817 A Taxon Fraxinus tomentosa pumpkin ash Fraxinus tomentosa pumpkin ash Quercus x bebbiana Bebb oak Quercus shumardii var. schneckii variant of Shumard oak Tilia americana American linden Gleditsia triacanthos honeylocust Grid 27SW 27SW 25Se (meters\/feet) Height (centimeters\/inches) DBH Year accessioned 1929 1929 1916 24.1 \/ 79.1 25.1 \/ 82.3 17.4 \/ 57.1 82.2 \/ 32.4 58.5 \/ 23.0 29.5 \/11.6 16883 A 32nW 21.3 \/ 69.9 61.7 \/ 24.3 1916 19804 A 21588 A 7Se 21ne 14.2 \/46.6 18.4 \/ 60.4 76.1 \/ 30.0 56.4 \/ 22.2 1916 1929 a period of growth and the laying of groundwork. now began in earnest his tireless, distinguished journey into science . . . In the decade from 1905, when he reorganized and restarted his Indiana herbarium and numbering system, through 1914, his last full year without a car, he averaged collecting about 1,500 specimens a year. But in 1915 alone he added 3,764\" (Kriebel 1987). Deam sent his collections to the Missouri and new york Botanical gardens and to Charles S. Sargent, director of the Arnold Arboretum. he asked for assistance in identifying his specimens. The Sargent Letter Books, found in the archives of the Arnold Arboretum, contain copies of thirty letters written by Sargent to Deam during the years 1914 through 1919. They reveal that Sargent identified over 600 tree and shrub specimens mounted on Deam's herbarium sheets. Sargent thought highly of Deam's work, writing on two occasions in 1915: \"I am very pleased indeed with your collection [Cornus and Salix] and I think you have done a capital piece of work, and certainly you are adding greatly to the knowledge and distribution of Indiana trees\" and \"There is nothing in your Carya collection which I should not have expected from Indiana. It is a remarkably fine collection and of very great assistance to me.\" When Sargent believed one of the trees found by Deam would enrich the Arnold Arboretum's living collection, he requested Deam send seed for propagation. The table above lists some specimens grown from seeds sent by Deam that still survive in the Arboretum's living collection. the deam oak In Wells County, Indiana, about three miles northwest of Bluffton, stands an oak tree which is well into its second century of growth. Specimens from this tree were first collected on October 4, 1904, by Bruce Williamson, a young zoologist, and his father. The specimens were taken to Deam who forwarded them to professor William Trelease of the Missouri Botanical garden for identification. growing in proximity to this tree were many white (Quercus alba) and chinquapin (Q. muehlenbergii) oaks. Though reminiscent of Q. alba, the leaves were not as deeply lobed and its acorns were not as large as those of a white oak. 18 Arnoldia 69\/4 the arboretum's First deam oak In 1908, the arnold arboretum received two plants of Quercus muehlenbergii 20 Arnoldia 69\/4 geORge hIBBen Quercus x deamii 21 WeLLS COunTy gIS geORge hIBBen An aerial view of the the Deam Oak Monument Forest (upper left), the smallest preserve in Indiana. DOugLAS SunDLIng One of the grafted Deam oaks growing at the Arboretum's Dana Greenhouses. Bibliography Deam, C. C. 1912. Trees of Indiana. (published in the Indiana Board of Forestry Report for 1911, Bulletin no. 1, pp. 86"},{"has_event_date":0,"type":"arnoldia","title":"Book Excerpt: Writing the Garden: A Literary Conversation Across Two Centuries","article_sequence":3,"start_page":22,"end_page":29,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25531","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270a726.jpg","volume":69,"issue_number":4,"year":2012,"series":null,"season":null,"authors":"Rogers, Elizabeth Barlow","article_content":"Book ExcErpt: Writing the Garden: A Literary Conversation Across Two Centuries Elizabeth Barlow Rogers David R. Godine, Publisher, Jaffrey, New Hampshire, with funding from the Foundation for Landscape Studies and the New York Society Library. 2011. 312 pages. ISBN: 978-1-567902-440-4. Editor's NotE: It'S SPRING, and those of us who love to garden are happily sinking our fingers into the warming soil as we plant seeds, pull early weeds, and ruthlessly hunt down lurking cutworms. But after a hard day in the garden it's time to relax with a good book, and what better than a book about some of the best garden writers (or writing gardeners) of the past couple of centuries. In Writing the Garden, author Elizabeth Barlow Rogers presents insightful essays on the works of a diverse group of writers. Some are well known, others less so, but in their writing all present fascinating opinions about the nature of gardening and a deep love for the subject. Rogers groups the authors into sections based on their interests and importance to garden literature, such as \"Women in the Garden,\" \"travelers in the Garden,\" and the delightful \"Humorists in the Garden\" (it turns out I'm not the only gardener who goes slug hunting at night with a flashlight). In the following excerpt, \"Warriors in the Garden,\" we are reminded that the seemingly gentle art of gardening is full of highly opinionated practitioners. Writing the Garden 23 G ardening is nothing less than warfare with nature. With no respect for the cabbage or the rose, nature sends in her legions of hungry insects and foraging animals to wreak havoc. But there is another kind of warfare in the garden, one that is waged against fellow gardeners rather than garden pests. In this kind of warfare garden theory is often presented as a polemical diatribe against previous practices or contrary philosophies. For the reader, it is both instructive and amusing to argue or agree with certain opinionated writers and to refight the horticultural battles of yesteryear as they promulgate their passionate beliefs and ideas. \"Warriors in the Garden\" William Robinson If [Gertrude] Jekyll was the authoritative mother of a more naturalistic English garden style, her friend William Robinson (1838 24 Arnoldia 69\/4 Writing the Garden 25 It would be a mistake, as Robinson is at pains to point out, to assume that the wild garden is the same thing as the native-plant garden. It should, to the contrary, be considered an opportunity to naturalize the flora of other countries, for as he tells us: Naturally our woods and wilds have no little loveliness in spring; we have here and there the Lily of the Valley and the Snowdrop, and everywhere the Primrose and Cowslip; the Bluebell and the Foxglove take possession of whole woods; but, with all our treasures in this way, we have no attractions in or near our gardens compared with what it is within our power to create. there are many countries, with winters colder than our own, that have a rich flora; and by choosing the hardiest exotics and planting them without the garden, we may form garden pictures. Here it is important to pause a moment and consider again the term \"garden pictures,\" since it is so frequently found in the writing of both Robinson and Gertrude Jekyll. For these writers, garden pictures did not imply the same thing as the Picturesque, the term commonly used to describe the earlier garden style in which designed landscapes were created in accordance with the principles of landscape painting. the garden pictures they had in mind are perhaps better characterized as vignettes, small scenes of beauty that the eye takes in as discrete discoveries rather than as panoramic scenery. Jekyll's carefully positioned camera framed many charming, seasonal vignettes within Munstead Wood, and in The Wild Garden, Alfred Parsons's engravings give graphic expression to Robinson's words, which are never themselves lacking in descriptive power. this does not mean, however, that such garden pictures, whether verbal or illustrational, should be considered as so many floral incidents independent 26 Arnoldia 69\/4 Writing the Garden 27 issue with Blomfield's recommendations for a return to formality, and here it is Blomfield's turn to aim a few angry verbal arrows at Robinson. Heatedly, he rebuts Robinson's sarcastic barbs, accusing him of willful misinterpretation and ignorance of garden making as a form of art: Mr. Robinson neither gives us the definition, nor shows us where the art is or what it consists of. the trees are beautiful, and so are the flowers, but where is Mr. Robinson's art? What does it do for us, or for the trees or the flowers? His skill as a tree-planter, or as a flower-grower, is no doubt great, but that does not make him an artist, and by no possible wrestling of the term can he be called so on this ground only. Blomfield maintained, \"the formal treatment of gardens ought, perhaps, to be called the architectural treatment of gardens, for it consists in the extension of the principles of design which govern the house to the grounds which surround it.\" Discriminating between the two views of gardening--the formal and the naturalistic--he argues: the formal school insists upon design; the house and the grounds should be designed together and in relation to each other; no attempt should be made to conceal the design of the garden, there being no reason for doing so, but the bounding lines, whether it is the garden wall or the lines of paths and parterres, should be shown frankly and unreservedly, and the garden treated specifically as an enclosed space to be laid out exactly as the designer pleases. He strongly refutes the notion that the landscape gardener has a monopoly on nature: the clipped yew-tree is as much a part of nature--that is, subject to natural laws--as a forest oak; but the landscapist, by appealing to associations which surround the personification of nature, holds the clipped yew-tree to obloquy as something against nature. Again \"nature\" is said to prefer a curved line to a straight, and it is thence inferred that all the lines in a garden, and especially paths, should be curved. Now as a matter of fact in nature--that is, in the visible phenomena of the earth's surface--there are no lines at all; \"a line\" is simply an abstraction which conveniently expresses the direction of a succession of objects which may be either straight or curved. \"Nature\" has nothing to do with either straight lines or curved; it is simply begging the question to lay it down as an axiom that curved lines are more \"natural\" than straight. For Blomfield, it was not the Italian style of formal gardening that was instructive for contemporary gardeners; rather it was the old gardens of England that had not succumbed to the fashion for Baroque ornamentation or, 28 Arnoldia 69\/4 Writing the Garden 29 subsequently, the Picturesque. Nor did formality imply a great expanse as in the French garden, for \"some of the best examples of [the English garden] are on a comparatively small scale.\" However, Blomfield does not merely sing the praises of old English formal gardens. With an architect's eye for composition and detail, he criticizes these as well as the later gardens designed in the Picturesque style, his principal objects of censure. He maintains that the white marble statues of Bacchus and Flora at Wilton were a mistake: \"to attain its full effect [marble] wants strong sunlight, a clear dry light, and a cloudless sky. In the soft light and nebulous atmosphere of the north marble looks forlorn and out of place.\" An integrated overall plan is what counts most, so in discussing public parks he comes down hard on \"the spasmodic futility\" of Battersea Park where, without a dominant idea controlling the general scheme, \"merely to introduce so many statues or plaster casts is to begin at the wrong end. these are the accidents of the system, not the system itself.\" Blomfield is united with Robinson, however unintentionally, in despising the Gardenesque style and the gardener who would have the specimen dahlia banish the hollyhock and other simple, old-fashioned flowers. He equally hates plants in beds that \"make the lawn hideous with patches of brilliant red varied by streaks of purple blue.\" taking sarcastic aim at the Victorian head gardener, he asks, \"Would he plant them in patterns of stars and lozenges and tadpoles? Would he border them with paths of asphalt? Would he not rather fill his borders with every kind of beautiful flower that he might delight in, and set them off with grass and pleasant green?\" In Blomfield's mind, the desired relationship between the architect and the horticulturist should not end in a standoff, nor would it, if their responsibilities were divided thusly: \"the designer, whether professional or amateur, should lay down the main lines and deal with the garden as a whole, but the execution, such as the best method of forming beds, laying turf, planting trees, and pruning hedges, should be left to the gardener, whose proper business it is.\" In this regard, it is worth noting that Gertrude Jekyll achieved some of her most notable gardens in collaboration with the architect Edwin Lutyens. their sympathetic marriage of brick terracing and hedge-enclosed garden spaces created an Arts and Crafts landscape idiom that influenced Vita Sackville-West and Harold Nicolson at Sissinghurst and many other gardeners up to the present day. Providing an architectural frame uniting house and garden and giving structure to seasonal borders of sophisticated horticultural artistry, this type of design might be viewed as a synthesis of Robinson and Blomfield. the harmonizing of their opposing but ultimately complementary theories resulted in a style that made a virtue of formal structure as a foil for loosely composed \"garden pictures.\" In this way these important late-nineteenth-century garden writers can be said to have assisted in the redirection of English garden style at a critical time when vast estate grounds were beginning to become a thing of the past. Elizabeth Barlow Rogers is a writer on the history of landscape design and the cultural meaning of place. She is the president of the Foundation for Landscape Studies and was the founding president of the Central Park Conservancy. Writing the Garden recently won a 2012 Book Award from the American Horticultural Society. Note: the images that accompany this excerpt are engravings by Alfred Parsons from William Robinson's The Wild Garden, 1881 edition. "},{"has_event_date":0,"type":"arnoldia","title":"2011 Weather Summary","article_sequence":4,"start_page":30,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25530","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270a36d.jpg","volume":69,"issue_number":4,"year":2012,"series":null,"season":null,"authors":"Famiglietti, Bob","article_content":"2011 Weather Summary Bob Famiglietti 2 011 continued the trend of warmer than normal temperatures and above average precipitation that started in 2008. Plentiful moisture plus a long growing season allowed the Arboretum's plants to attain optimum growth. Some of our plants suffered damage from storms during the year. January began mild, and Arboretum visitors celebrated New Year's Day at 59 Weather 31 Arnold Arboretum Weather Station Data 32 Arnoldia 69\/4 Weather 33 8th and 9th. it never reached 90 34 Arnoldia 69\/4 Weather 35 Two inches of heavy, wet snow fell on October 30, 2011. The next day, traces of snow along with fallen leaves remained around this ginkgo (Ginkgo biloba, accession 222-97-a). DeceMber was very mild, continuing November's warmth. A high of 62"},{"has_event_date":0,"type":"arnoldia","title":"Wilson's Pearlbush (Exochorda giraldii var. wilsonii): A Gem to the Core","article_sequence":5,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25534","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270af28.jpg","volume":69,"issue_number":4,"year":2012,"series":null,"season":null,"authors":"Schneider, Stephen","article_content":"Wilson's Pearlbush (Exochorda giraldii var. wilsonii): A Gem to the Core Stephen Schneider I t can be a memorable experience the first time you crack open a geode--pale gray and nondescript on the outside, the colorful crystalline center is anything but. The same can be said for cutting into the wood of the trees and shrubs in the Living Collection at the Arnold Arboretum. There have been many surprises for Arboretum staff who prune and remove trees and are also interested in woodworking; often what is hidden by thick, scaly, neutral-colored bark proves to be a treasure once the inner wood is revealed. Several species come to mind when considering unique and beautiful wood. Golden raintree (Koelreuteria paniculata) and Osage orange (Maclura pomifera) yield consistent chocolate brown and bright yellow heartwood, respectively. Boxelder (Acer negundo), on the other hand, often displays an erratic, bright red fungal staining in parts of its center. Even the oldgrowth stems of common lilac (Syringa vulgaris) often have a deep purple center that, unfortunately, disappears once the wood is seasoned. Although there are many other Arboretum plants that possess interesting wood, a large specimen of Wilson's pearlbush (Exochorda giraldii var. wilsonii, accession 11626-C) merits particular attention. Grown from seeds collected in 1907 by E. H. Wilson in Hubei, China, this centenarian shrub resides just off the road near the top of Bussey Hill. Its racemes of spring flowers start as white, pearllike buds and open to perfect, five-petaled flowers. The flowers are followed by interesting star-shaped seed capsules. Mature and well established, this multi-stemmed shrub has a commanding spread of about twenty feet and a height to match. Its presence, however, is often overlooked by the many visitors who pass by it each day on their march to the top of the hill. They are unaware of the secret that lies beneath its bark. I remember well the first time I was introduced to Exochorda wood. A rather small piece, about a foot long and four inches in diameter, was tossed to me from across the room. Its weight took me by surprise--it felt as strong and dense as hickory. A first attempt to cut through it failed, since the wood was too hard for the band saw blade to provide a straight cut. It became necessary to use a fine-toothed carbide blade on a table saw. That machine even seemed to struggle a bit, but the results were worth the effort. Hidden beneath the gray, scaly, exfoliating bark was densely grained wood patterned in light and dark browns with orange-red highlights throughout. A single pass of the blade proved to be all that was needed to create a smooth finish, velvety to the touch. Applying a coat or two of Danish oil enhances the beauty of this material since it makes the swirling grain more noticeable. When put on a lathe and turned, this wood creates a beautiful spindle that displays the variety and complexity of its colors and patterns. Checking (cracking that occurs during the lumber drying process) is nearly impossible to avoid with a wood this dense, so finding stable stock to work with between the cracks can be a challenge. Since discovering the wood of Exochorda, I have reserved the use of it for very special projects for very special people. Since Wilson's pearlbush is a relatively easy plant to grow, I'll often give the recipient of the gift a live specimen of it to plant in the backyard as a reminder that, much like a geode, its plain appearance on the outside can harbor profound beauty on the inside. Stephen Schneider is Manager of Horticulture at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23428","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd1608528.jpg","title":"2012-69-4","volume":69,"issue_number":4,"year":2012,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Picking Up the Pawpaws: The Rare Woody Plants of Ontario Program at the University of Guelph Arboretum","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25528","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260896f.jpg","volume":69,"issue_number":3,"year":2012,"series":null,"season":null,"authors":"Fox, Sean","article_content":"Picking Up the Pawpaws: The Rare Woody Plants of Ontario Program at the University of Guelph Arboretum Sean Fox t might surprise you to learn that, in Canada, species such as Magnolia acuminata (cucumbertree), Betula lenta (sweet birch), and Morus rubra (red mulberry) are among those that are listed as endangered in the wild (see Table 1). You may be thinking, \"Really, Betula lenta? It grows all over the place in the eastern United States!\" It's true that the majority of woody plant species at risk in Canada are quite secure in the United States, so why the concern? Is there really a need for conservation? Conserving an organism at the species level is generally regarded as the most immediate and crucial objective of many conservation programs. In the case of a species that is critically endangered on a global scale, simply ensuring the survival of a few individuals is often a significant challenge. But even many globally common plant species have conservation needs. Often these species do not have a very high representation of diverse genetic material archived in ex-situ collections simply because they are not considered to be a high priority for conservation. To compound this, the limited germplasm that is archived is often accessed from similar populations from the core of a species geographic range. By collecting from more provenances, including those at the extremes of a species' range, we can come closer to fully conserving and representing the genetic diversity of the species. After the Laurentide Ice Sheet began receding nearly 12,500 years ago, the forests of eastern North America began their march northward. Species migration is a dynamic and ongoing process, and while many species have already pushed into the tundra region in the far north of Canada, most other species have only extended into southern Canada far more recently. These regional populations, on the forefront of a long migration into northern latitudes, must adapt I to an array of environmental conditions that are often very different from those found at the core of the geographic range. Adapted gene complexes enable a plant to adjust to the timing of the local annual growth cycle, including bud break, root growth, shoot and leaf elongation, bud development, diameter growth, and cold acclimation. The genetic variation present in NANCY ROSe Notable for its bright yellow fall foliage, sweet birch (Betula lenta) is a rare find in Canada. The Rare Woody Plants of Ontario Program 3 ALL PHOTOS BY THe AUTHOR UNLeSS OTHeRWISe INdICATed Flowering dogwood (Cornus florida) blooming in Ontario. these range extensions is very significant from a conservation standpoint since these particular genotypes may provide crucial genetic material to allow a species to migrate and fill various regional niches. The Ontario populations of woody species, at the northern extent of their natural range, represent adaptations to our northern conditions. Liriodendron tulipifera from Ontario are more likely to be suitable for forestry planting in that province than seedling stock from a Virginia source. Cornus florida from Ontario-based provenances have proven, in cold hardiness trials, to be more winter hardy in Canada than nursery stock sourced from farther south. As migration pressures increase due to a rapidly changing climate, it may become even more critical to conserve these northern genotypes. Unfortunately, the pace of abiotic change in the environment is likely to be far ahead of biotic survival for many species. The continued exploitation and segregation of suitable habitat adds another dynamic to an already challenging scenario for in-situ conservation. UndeRsTAndinG sPecies AT Risk in cAnAdA Taking the time to thoroughly understand the legislative conditions regulating species at risk in Canada, as in many other parts of the world, can be an exercise in patience. The federal government of Canada's National Strategy for the Protection of Species at Risk is composed of the National Accord for the Protection of Species at Risk (NAPSR), established in 1996; the Habitat Stewardship Program (HSP), established in 2000; and the Species at Risk Act (SARA) established in 2003. The National General Status Working Group (NGSWG) was formed in 1996 to support the mandate of the NAPSR, and is charged with establishing status rankings for all species in 4 Arnoldia 69\/3 cucumbertree (Magnolia acuminata): canada's First endangered Tree Magnolia acuminata was the first tree in Canada to be listed as endangered by COSeWIC in 1984. In 2003, this species was re-evaluated as endangered under the SARA and plans for a recovery strategy were developed. Cucumbertree has always had a very limited distribution in Canada and is currently only known to occur in two areas of southwestern Ontario. In total, only 283 naturally occurring individuals are known to survive in 12 small, extant populations. These individuals represent an extreme northern extension for a species that forms its most abundant core population in the central Appalachian range of the United States. The cucumbertrees remaining in Ontario are generally in good health; however, the habitat that supports them is highly fragmented. This segregation has not only reduced the reproductive fitness of the remaining populations (perhaps due in part to a reduction in pollinator-supporting habitat), but it has also eliminated suitable conditions for seedling regeneration. The range of cucumbertree also happens to lie within the most heavily populated area of human settlement in Canada and one of the most rapidly-developing regions in North America. In-situ conservation efforts to identify and protect individual trees in isolated woodlots have had some success. However, further steps are required to ecologically connect these remaining sites in order to allow this magnificent species to continue its natural migration within Ontario. cucumber Tree Magnolia acuminata N species Found non-Planted > 100 species Found non-Planted 20 6 Arnoldia 69\/3 The Rare Woody Plants of Ontario Program 7 PeGGY HURST A series of interpretive plaques were created for Ontario's rare woody plants with support from BGCI Canada's Investing in Nature: A Partnership for Plants program. Here, Kentucky coffeetree (Gymnocladus dioicus) is highlighted in the University of Guelph Arboretum's World of Trees collection. John Ambrose (right), with botanists Lindsay Roger and Gerry Waldron, upon their discovery of a new species to Canada, swamp cottonwood (Populus heterophylla), in 2002. The eastern Redbud (Cercis canadensis): O canada--its home and native land? A specific epithet like \"canadensis\" might lead one to believe that eastern redbud floods the understory of the great northern forests. But, despite its seeming patriotism to Canada, this beautiful species is not quite as common in the north as one might think. In Gerry Waldron's wonderful book, Trees of the Carolinian Forest (2003), he quotes the great Canadian botanist, John Macoun, as he recounts his first and only sighting of eastern redbud on Canadian soil in 1892: ... I was informed that a remarkable tree grew on the south end of the island, that many years ago produced an abundance of lovely red flowers in early spring before the leaves came out ... the next day I examined the south point and found the tree. It had been undermined by the waves and fallen inland, and more than half its limbs were dead, but it still bore leaves and what remained was quite healthy. It will soon disappear, but the record of its existence will remain. These flower buds of eastern redbud (Cercis canadensis) show the species' interesting trait of cauliflory (flower and fruit production from woody stems). This accession (1988-0284.002) in the World of Trees collection at the University of Guelph Arboretum is from a cold-hardy provenance in Wayne County, Michigan. This tree that Macoun happened across remains the only naturally-occurring individual ever discovered in Canada. This plant grew at Fish Point, Pelee Island, in Lake erie--the most southerly point in all of Canada, and, as he predicted, was eventually swallowed by the lake as the shoreline eroded away. While there are naturalized populations established in parts of southwestern Ontario, as escapees from cultivated stock, eastern redbud is now officially ranked as extirpated in Canada. 8 Arnoldia 69\/3 ROB GUTHRIe The Rare Woody Plants of Ontario Program 9 Table 2. Accessions of known, wild, Ontario-based provenance for selected rare woody taxa under cultivation at the University of Guelph Arboretum. Taxon Aesculus glabra Asimina triloba Betula lenta Campsis radicans Carya laciniosa Carya glabra Castanea dentata Celtis tenuifolia Cornus drummondii Cornus florida Euonymus atropurpurea Fraxinus profunda Fraxinus quadrangulata Gleditsia triacanthos Gymnocladus dioicus Hibiscus moscheutos Juglans cinerea Liriodendron tulipifera Magnolia acuminata Morus rubra Morella pensylvanica Pinus rigida Platanus occidentalis Ptelea trifoliata Quercus ellipsoidalis Quercus muehlenbergii Quercus prinoides Quercus shumardii Rosa setigera * G-global, S-provincial G1-extremely rare, G2-very rare, G3-rare to uncommon, G4-common, G5-very common S1-critically imperiled, S2-imperiled, S3-vulnerable, S4-apparently secure, S5-secure Risk Ranking* G5, S1 G5, S3 G5, S1 G5, S2 G5, S3 G5, S3 G4, S3 G5, S2 G5, S4 G5, S2 G5, S3 G4, S2 G5, S3 G5, S2 G5, S2 G5, S3 G4, S3 G5, S4 G5, S2 G5, S2 G5, S1 G5, S2 G5, S4 G5, S3 G5, S3 G5, S4 G5, S2 G5, S3 G5, S3 Total Number of Accessions 5 8 9 3 6 3 2 5 5 8 6 1 20 7 26 1 12 11 16 5 3 4 10 22 2 16 2 4 6 Total Number of Individuals 20 12 44 4 25 7 3 13 26 17 16 3 26 38 87 2 32 15 37 21 3 5 18 43 2 64 9 9 8 10 Arnoldia 69\/3 kentucky coffeetree (Gymnocladus dioicus): distribution within the University of Guelph Arboretum WHILe gene banking various accessions within seed orchards makes archiving and maintaining plant material simpler, a strong effort has also been made to establish accessions in suitable botanical and horticultural collections throughout the Arboretum. distributing our conservation collections in this fashion serves several purposes: 12 Arnoldia 69\/3 The Rare Woody Plants of Ontario Program 13 with a particular emphasis on recently discovered species such as Quercus ellipsoidalis (1978), Fraxinus profunda (1992), Quercus ilicifolia (1994) and Populus heterophylla (2002). These are important species that we hope to further incorporate into our ex-situ collections at the Arboretum. As our existing seed orchards continue to produce increasingly sound crops, we are now in the position to better distribute this seed to nurseries and local conservation authorities to aid in their restoration activities. Large crops of seed will also be archived at the National Tree Seed Center in Fredericton, New Brunswick, and the Ontario Tree Seed Plant in Angus, Ontario. Seed will continue to be available to other botanical institutions for conservation and research purposes. In this modern era, and with an unstable economy, most botanical gardens and arboreta are facing tough challenges with budget and staff cuts. As the years have progressed at the University of Guelph Arboretum, we've also had to make difficult decisions regarding the activities that we have the capacity to engage in successfully. While we've had to scale back several of our display-based horticultural collections, we've found that our conservation programs have helped to provide a niche that further defines the mission of our organization. It must always be remembered that ex-situ conservation programs, as valid and critical as they are, don't hold a candle to ecosystem conservation, expansion, and linkage. These in-situ conservation activities must be represented in our highest aspirations as citizens and nations. However, the important role that botanical gardens and arboreta can play must not be underestimated either. Whether it is the education, outreach, research, stewardship, or conservation hat that is being worn, public gardens are in a unique position to be meaningfully engaged in rare flora programs both locally and globally. References Ambrose, J. and d. Kirk. 2006. Recovery Strategy for Cucumber Tree (Magnolia acuminata L.) in Canada. Prepared for the Ontario Ministry of Natural Resources by the Cucumber Tree Recovery Team, Toronto, Ontario. Beardmore, T., J. Loo, B. McAfee, C. Malouin, and d. Simpson. 2006. A Survey of Tree Species of Concern in Canada: the Role of Genetic Conservation. The Forestry Chronicle 82-3: 351"},{"has_event_date":0,"type":"arnoldia","title":"Land Bridge Travelers of the Tertiary: The Eastern Asian-Eastern North American Floristic Disjunction","article_sequence":2,"start_page":14,"end_page":23,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25526","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d2608526.jpg","volume":69,"issue_number":3,"year":2012,"series":null,"season":null,"authors":"Yih, David","article_content":"Land Bridge Travelers of the Tertiary: The Eastern Asian The Eastern Asian 16 Arnoldia 69\/3 18 Arnoldia 69\/3 MIcHAEl DOSMANN 20 Arnoldia 69\/3 22 Arnoldia 69\/3 MIcHAEl DOSMANN "},{"has_event_date":0,"type":"arnoldia","title":"A Rare Find: Yellow-Fruited Spicebush (Lindera benzoin forma xanthocarpum)","article_sequence":3,"start_page":24,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25524","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d2608128.jpg","volume":69,"issue_number":3,"year":2012,"series":null,"season":null,"authors":"Lynch, Richard","article_content":"A Rare Find: Yellow-Fruited Spicebush (Lindera benzoin forma xanthocarpum) Richard Lynch here are as many different harbingers of spring as there are fond memories in the minds of the people who look for them. For some, the last of the snow melting off a northern slope fits the bill. For others, the first chorusing of spring peepers (Hyla crucifer) in the still-cold ponds of late March provides hope for the warmer seasons to come. For those with a more botanical bent, and especially for lovers of the deep woods across the eastern United States, the opening of the tiny yellow flowers of spicebush (Lindera benzoin) clearly marks the tipping point from winter to spring. In many parts of the Northeast, spicebush is the first shrub to flower and is often timed with the arrival of mourning cloaks (Nymphalis antiopa) and spring azure butterflies (Celastrina ladon). Spicebush also plays a role in alerting nature lovers that the fall season approaches. By the middle of September, female plants begin to display some of the brightest red fruit found in nature. Plants growing in deep woods will be a bit sparse in fruit, but those growing along the forest edge or near wetlands can produce a great profusion of colorful fruit. There are a great number of resident and migrating bird species that take full advantage of the bounty, and often within a week or two the T The typical bright red fruit of spicebush (Lindera benzoin). all PhoToS By The aUThoR UNleSS oTheRwISe INdIcaTed Lindera benzoin forma xanthocarpum bears golden-yellow fruit. NaNcy RoSe Yellow-Fruited Spicebush 25 fruit have been harvested and carried off by wildlife. Though bright red is the typical fruit color, there is also a yellow-fruited spicebush (Lindera benzoin forma xanthocarpum). The story of this unusual variant begins in Shrewsbury, Massachusetts, in 1913, where it was discovered by Mrs. Frank e. lowe. a description of the plant written by G. S. Torrey was published the following year in Rhodora (note that the species name was then Benzoin aestivale rather than the current Lindera benzoin): \"on october 4, 1913, Mrs. Frank e. lowe collected in Shrewsbury, Mass., specimens of the Spice Bush, Benzoin aestivale Nees., which differed from the common form in having the drupes orange-yellow, instead of bright red. Several bushes were found, some growing with the typical form in low, damp places; some alone, in drier ground in a rocky pasture. They all bore yellow fruit, which were ripe and falling. The material was sent by Mrs. lowe to Mrs. e. l. horr of the worcester Natural history Museum, by whom it was referred to the Gray herbarium. The plant may be characterized as follows: BeNZoIN aeSTIVale (l.) Nees., forma xanthocarpum, forma nova, fructus flavis.\" A view across the lowland sweetgum 26 Arnoldia 69\/3 Yellow-Fruited Spicebush 27 Green milkweed (Asclepias viridiflora), left, and globose flatsedge (Cyperus echinatus), right, are two of the rare (in New York) plants found growing in a sunny meadow within the Staten Island Greenbelt. rubrum), where a few of the golden-yellow fruit still remained. after the discovery of the first specimen, we made a greater effort to look for the yellow-fruited plants in the vicinity of the first one, but found none. Further up the trail and into a sunny meadow, we discovered two more plants heavily laden with yellow fruit. The meadow is part of a successional grassland growing over serpentine-derived soils and contains other New york State rare plants such as green milkweed (Asclepias viridiflora), purple milkweed (A. purpurea), and globose flatsedge (Cyperus echinatus). we collected these additional yellow fruit and added the fruit from the first collection; these became a separate cohort of seed from which we could propagate. It turns out that the yellow-fruited spicebush is a rarer taxon than first believed. according to charles Sheviak, the state botanist for New york, the plant had not previously been recorded as growing in the state. In Massachusetts, state botanist Brian connelly has no records for any extant populations in the state. although it is likely that other populations for the plant do exist, there are no other confirmed populations known in these two states. The question then arises as to what conservation efforts, if any, need to be taken to ensure the continued existence of the plant in the wild. In using the term \"forma\" in describing the plant, G. S. Torrey seems to convey the belief that the plant is a random mutation, not sustainable over time, occurring within a larger population. In general, the term \"variety\" would be used to describe a plant that is self-sustaining or represents a variant that covers a portion of the range of a larger species description. Given that we don't know either the genetic differences that separate the yellow-fruited spicebush from the more common red-fruited 28 Arnoldia 69\/3 "},{"has_event_date":0,"type":"arnoldia","title":"Book Review: A Landscape History of New England","article_sequence":4,"start_page":29,"end_page":34,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25525","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260816d.jpg","volume":69,"issue_number":3,"year":2012,"series":null,"season":null,"authors":"Andersen, Phyllis","article_content":"Book Review: Phyllis Andersen A Landscape History of New England Edited by Blake Harrison and Richard W. Judd. Cambridge: MIT Press, 2011. 413 pages. ISBN 978-0-262-01640-7 cultivation by Native Americans was ignored in the jeremiads of early Puritan ministers who needed a transformative narrative to motivate their flocks to both stay and spread out. As waves of settlers came to understand the intrinsic capacities of the landscape, the wilderness became a land of cultivation and harvesting: pastures, orchards, and gardens; forests for fuel and building material; rocky and sandy coastal waters offering access to a rich diversity of fish and crustaceans. Joseph Conforti opens the roster of essays by setting a theme for the entire volume: regional identity as both historically grounded and culturally invented. Conforti projects New England identity as flowing from Native American C oeditors Blake Harrison and Richard W. Judd challenged a group of senior and young scholars to produce essays that capture myriad aspects of the New England landscape: the material landscape of forests, upland farms, stone walls, inland rivers, and rocky coast lines, and the symbolic landscape of picturesque villages, bucolic pastures, and the stock pieties of hard-working farmers with backs to the plow and eyes on the horizon. Methodologies deployed by the authors vary from the new disciplines of environmental and ecological history to literary narrative and to the politics of gender, ethnicity, and environmental change. The twenty essays are bookended by the editors' introduction and conclusion--dissimilar threads skillfully woven to form comprehensive case studies of landscape and cultural changes over three centuries. The essays engage both the essence of regional character and the theatrical promotion of magnetic scenery created for the seduction of tourists to visit New England and support local economies. Old England was a refuge for New England's early settlers, so newly settled places were often named after mother-country places (the Berkshires, Portsmouth, Worcester, New London) and topographic terms (brook, pond, marsh, fens) coincidental to mother country terms. This offered familiarity amidst what some early settlers called the emptiness of the place and others called the howling wilderness. The fact that the \"emptiness\" contained areas of NEW HAMPSHIRE HISTORICAL SOCIETy 30 Arnoldia 69\/3 A Landscape History of New England 31 \"Fishermen and weir, Passamaquoddy Bay region near Eastport, Maine, circa 1880. This photograph was part of broader study by the U.S. Fish Commission for the 1880 U.S. Census. It shows fishermen using a traditional brush weir to take herring for the burgeoning canning industry.\" tribes such as the Algonquians, with their seasonal settlements and cultivation of crops, to the formation of isolated towns and villages distributed across farmland and along the seacoast, a land-planning method still visible today. The New England landscape was physically reconstituted in the nineteenth century with a surge of industry, especially shoe manufacturing, textile mills, and ship building. The current evolutionary stage of development includes a topology of leisure and recreation: heritage sites, boutique-lined waterfronts, ski slopes, athletic fields, and the indigenous clothier of fishers and hunters, L. L. Bean. Conforti quotes Dona Brown, a historian at the University of Vermont, to describe that tourist landscape as \"a commodity peddled and consumed like the notions of an itinerant yankee trader.\" In his essay, Kent Ryden finds the well-worn argument of nature vs. culture a useless bit of rhetoric in understanding the New England landscape. Everything we see is the result of land use, he insists, recorded in the ways that human minds and hands worked in tandem with natural opportunities and constraints. He cites a little-known essay by Thoreau, \"The Succession of Forest Trees,\" first delivered as a lecture in 1860. From years of observing \"Tourists in Franconia Notch, 1920s. Franconia Notch was one of the most popular sites in New Hampshire's White Mountains. Here, tourists by the shores of Profile Lake gaze upward at the Old Man of the Mountain.\" Source: From Automobile Blue Book. T. W. SMILLIE, NATIONAL ARCHIVES PHOTO 22-CD-451 32 Arnoldia 69\/3 LIBRARy OF CONgRESS A Landscape History of New England 33 control. Attracted to a life of self-sufficiency, writers in particular were drawn to the back country of New England where they documented their survival tactics in numerous publications. Dona Brown describes a little known back-to-the-land movement of the 1930s; she notes that an \"imaginative reconfiguration\" of New England was underway as the image of a region full of \"dour puritans and antiquated blue laws\" was refigured. In a 1932 editorial in Harper's Monthly, the noted writer and historian Bernard DeVoto observed that the great Depression was not as severe in New England because long years of trials and tribulations had given the people great moral strength: \"By the granite they have lived for three centuries, tightening their belts SPECIAL COLLECTIONS, RAyMOND H. FOgLER LIBRARy, UNIVERSITy OF MAINE \"Henry Red Eagle on the shores of Moosehead Lake, circa 1940. Red Eagle often drew inspiration from the Moosehead Lake region, incorporating its recreational and its working spaces into his writing.\" Source: From Bangor & Aroostock Railroad, In the Maine Woods (1941). and hanging on.\" Brown uses as an example the writer Elliott Merrick and his wife who gave up urban life for a back country farm in Vermont where Merrick wrote From This Hill Look Down (1934). He stressed self-reliance and hard work in taming nature as a way to revitalize the urbanized mind and body. The couple was followed by another pair of writers, Helen and Scott Nearing, who relocated first to rural Vermont and then Maine. Their book, Living the Good Life (1954), became a manual for disaffected youth of the 1960s and 1970s. Elizabeth Pillsbury investigates Long Island Sound on New England's southerly shore, valued first for its oysters and later for boating recreation. The Sound became a waste depository and ended up as a dead ecosystem. Moving up the coast line, Robert gee brings his reader to Maine's \"drowned coast\" created by the rising and then receding sea level revealing land features: dramatic inlets of eroded tide pools and island clusters accommodating a rich variety of sea and shorebird life. gee tracks the development of Maine's fish canning and blueberry industries in tandem with its growing popularity for tourists and summer homes. Moving back down to Boston, Michael Rawson traces the concern for the environmental health of Boston Harbor today back into the nineteenth century, when extensive filling of brackish tidal flats dramatically altered the shore line. The topic of alternative ways of writing about the New England landscape is covered by two essays on lesser known individuals, each dealing with the ambiguity between documentary and fantasy writings. Under the pen name Henry Red Eagle, the Native American writer and wilderness guide, Henry Perley, wrote numerous stories about Maine's north woods. Written for a popular audience, his tales of adventure and romance highlighted his Native culture. Perley also participated in tourist activities, and like many other Natives took roles in national performing troupes such as P. T. Barnum's, cooperating with displays of stereotypical Indian behavior demanded as entertainment by \"white man\" audiences. Similarly, the Maine travel writer george H. Haynes, who, in the words of contributing author David L. Richards, spe- 34 Arnoldia 69\/3 "},{"has_event_date":0,"type":"arnoldia","title":"Remember Last Winter?","article_sequence":5,"start_page":35,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25527","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260856b.jpg","volume":69,"issue_number":3,"year":2012,"series":null,"season":null,"authors":null,"article_content":"Remember last winter? MICHAEl DOSMANN This image of Meadow Road and the Fabaceae (legume family) collection was made on January 13, 2011, after yet another heavy snowfall. Read a summary of 2011 weather events at the Arboretum in the next issue of Arnoldia. 36673667 U.S. POSTAL SERVICE STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION (Required by 39 U.S.C. 3685) 1. Publication Title: Arnoldia. 2. Publication No: 0004"},{"has_event_date":0,"type":"arnoldia","title":"Plainly Unique: Schisandra chinensis","article_sequence":6,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25529","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d270a328.jpg","volume":69,"issue_number":3,"year":2012,"series":null,"season":null,"authors":"Schmerler, Sam","article_content":"Plainly Unique: Schisandra chinensis Sam Schmerler T he plants of the Arnold Arboretum display incredible floral diversity. Magnolia macrophylla's huge waxy blooms open twice, partly closing in between for an overnight sex change. Helwingia japonica sprouts tiny green umbels in the center of otherwise unremarkable leaves. Davidia involucrata forgoes petals entirely, but shelters its reproductive organs with massive white bracts. Even wild Viola sororia, flagging down bees with its iconic violets, surreptitiously sends out discrete, selfpollinating flowers underground. With all this bizarre and beautiful reproduction going on, most of us overlook the most evolutionarily distinctive flowering plant in the collection: Schisandra chinensis. An unassuming woody vine, it represents a unique and ancient lineage that parted ways with most other flowering plants at least as far back as the early Cretaceous, before even \"living fossils\" like Magnolia. This ancient group, the Austrobaileyales, is now recognized as the third-oldest remaining branch on the phylogenetic tree of flowering plants, diverging after only Amborella (a strange New Caledonian shrub) and the Nymphaeales (a group of herbaceous aquatics that includes water lilies). This means that all the other flowering plants in our collection--from creeping crowberries to towering tuliptrees--are more genetically similar to each other than any of them are to Schisandra. We can't grow the other Austrobaileyales here, since they hail from warmer forests in North America, Asia, and Oceania, but Schisandra chinensis, from temperate northeastern Asia (China, Korea, northern Japan, eastern Russia), can reliably survive Boston winters. This dioecious vine doesn't appear particularly primitive. Visually, it doesn't stand out much at all. Our two specimens (343-97-B, a male plant from Changbaishan, China, and 409-97-B, a female from Chiaksan, South Korea) twine unobtrusively up his-and-hers trellises in the Levintritt Shrub and Vine Garden and tend to blend in with their neighbors. Their simple, medium-green leaves are perfectly innocuous, eventually turning a bland butter yellow. In late spring, small, white, sweet-smelling flowers droop on thin pedicels in a passable impersonation of nearby Actinidia (kiwi). The female's flowers develop into elongated fruits with numerous bright red, berrylike fruitlets. Winter will reveal exfoliating bark resembling that of climbing hydrangea. Evolutionary biologists (including Arboretum director Ned Friedman) have discovered that Schisandra and the other Austrobaileyales can offer insight into many key events in the history of flowering plants. Aspects of Schisandra's vascular system may represent an early step in the development of vessels, the structures that allow most flowering plants to rapidly transport water and ecologically dominate hot and dry habitats. Schisandra also retains a relatively simple anatomy during its haploid stage, with only four nuclei and one developmental module in each female gametophyte (almost all flowering plants have eight nuclei and two modules). The endosperm of Schisandra seeds consequently contains only one complement of genes from each of its parents, while most flowering plants acquire an additional copy of their moms' genes. Schisandra likely shares these characteristics with the extinct ancestors of all flowering plants, a living link to the distant past. But while it retains many archaic anatomical features that are long lost in most flowering plants, Schisandra has evolved a unique and medically promising biochemistry. Traditional Chinese herbalism prescribes S. chinensis for a whole host of ailments and as a general tonic and adaptogen. Recent science has isolated several new types of lignans (a class of polyphenols) from the fruits; these have anti-oxidant and anti-inflammatory properties. Schisandra lignans have been shown to protect brain cells from glutamate and liver cells from a variety of toxins; they may also inhibit platelet aggregation, tumor proliferation, and possibly even HIV replication. As chemists in the food and medical industries increasingly explore these lignans, it's likely that demand for S. chinensis as a pharmaceutical precursor and \"functional food\" will increase. Even though Schisandra may not dazzle, this vine's exciting chemistry and singular evolutionary history prove it truly stands alone. Next time you visit the Arboretum be sure to check out Schisandra chinensis--it tends to reward closer inspection. Sam Schmerler recently completed his appointment as a Curatorial Fellow at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23427","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160816e.jpg","title":"2012-69-3","volume":69,"issue_number":3,"year":2012,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"One Hundred Years of Popular Information","article_sequence":1,"start_page":2,"end_page":10,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25521","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260b726.jpg","volume":69,"issue_number":2,"year":2011,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"one Hundred years of Popular information Peter Del Tredici Publications History O n May 2, 1911, the Arnold Arboretum published the first issue of the Bulletin of Popular Information. Arboretum director Charles Sprague Sargent was the editor, and he stated that its specific goal was to meet the complaints of people who \"... do not know when the trees and shrubs in the Arboretum bloom and therefore miss flowers which they want to see.\" These first bulletins came out once a week during the spring, early summer, and fall and were \"mailed without charge to anyone interested in trees and shrubs and their cultivation.\" They were four pages long and without illustrations. The intention of the publication was to tell people who lived in the area what exciting things were happening on the grounds of the Arboretum and to provide some basic facts about selected plants, including their history of cultivation and suitability for New England gardens. The fact that the Bulletin came out thirty-nine years after the establishment of the Arboretum in 1872 suggests that public outreach to visitors was not originally very high on Sargent's to do list. By 1911, however, he apparently felt that the time was ripe to connect with the gardening public who wanted to know more about the collections. Despite its tardy introduction, the Bulletin of Popular Information established an Arboretum tradition of outreach through publications that continues today. After four years of publication, Sargent initiated a \"New Series\" of the Bulletin on April 28, 1915. Unlike the first series, this one had volume numbers and an index and established a subscription rate of one dollar. The first issue of the second series concluded with the rather quaint note that, \"Automobiles are not admitted to the Arboretum, but visitors who desire carriages to meet them at the Forest Hills entrance can obtain them by telephoning to Charles S. Sargent, first director of the Arnold Arboretum, photographed at the Arboretum in 1904. P. J. Brady, Jamaica 670, or Malone & Keane, Jamaica 344.\" From the perspective of today's digitally connected world, it's hard to imagine a time of when entire telephone numbers rather than just area codes consisted of three digits. Series Three of the Bulletin began in April 1927--a month after Sargent's death--edited by Ernest H. Wilson. It was printed on coated paper for the first time, which allowed for the insertion of a full page black-and-white photograph in each issue. Following Wilson's untimely death in an automobile accident on October 15, 1930, long-time Arboretum staff member ArCHIvES OF THE ArNOlD ArBOrETUM Bulletin of Popular Information 3 J. G. Jack took over the task of producing the Bulletin with contributions from Oakes Ames, W. H. Judd, and the young Edgar Anderson. Anderson took over full responsibility for the publication in 1932 and initiated a fourth series in 1933. In 1935, Anderson left the Arboretum to work at the Missouri Botanical Garden and horticulturist Donald Wyman took over the Bulletin, publishing his first article on \"Tree Troubles\" in March 1936. Elmer Drew Merrill had been appointed director of the Arboretum in 1935 and in March 1941 he made the decision to change the name of the Bulletin to Arnoldia. He had two reasons for doing so: first, he thought that the title Bulletin of Popular Information was too cumbersome, and second, that it was difficult to cite in scientific papers because it had been published in four separate series without sequential volume numbers. He also felt that changing the name to Arnoldia would not only \"reflect proper institutional credit on its sponsoring institution.\" Merrill retained Wyman as editor of the newly christened publication, a post Wyman held until 1969--a 34-year record of longevity that no one is ever likely to top. During the entire period of Wyman's editorship, Arnoldia came out more or less twelve times per year with each issue being of variable length. Following Wyman's retirement in 1969, richard Howard, who served as director from 1954 through 1977, changed Arnoldia from its pamphlet format to a magazine format in 1970. A card-stock cover with a full-bleed (printed NANCy rOSE Bound copies of every issue of the Bulletin of Popular Information and Arnoldia are housed in the Arboretum's library. All issues are also available through the Arboretum's website (http:\/\/arnoldia.arboretum.harvard.edu). 4 Arnoldia 69\/2 Bulletin of Popular Information 5 hold its own in New England. The hardiness of Writers and content this tree, its rapid growth, and the fact that it It is worth noting that the editors of the publiis not injured by insects, suggest that this is a cation during its first fifty-eight years--Sargent, good subject to plant in narrow streets. Seeds Wilson, Anderson, and Wyman--were also its will be sent from the Arboretum in the autumn principle writers. This aspect gave it both a to anyone who may desire to grow this tree.\" highly personal and authoritative tone. The This fascinating quotation reveals much people who were writing about the plants knew what they were talking about and, because the about the early history of the Arboretum that, Arboretum was a research and not a commercial had it not been written down, would have been institution, they could be counted on for unbiased information. All of these botanists wrote with a high level of confidence based on the completeness of the Arboretum's collections and on the soundness of their observations and judgment. Issues of the Bulletin from the Sargent years, in particular, make for fascinating reading because they tell the story of the early introduction of many now-familiar plants. Sargent was also very adept at capturing the essence of plants, sometimes in completely unexpected ways, such as his description of Populus tomentosa leaves, which \"hang on long flattened stalks and, fluttering in the slightest breeze, make, as the blades come together, a noise like drops of rain in a heavy shower falling on a tin roof\" (July 2, 1915). Many of the plants that were first described in the Bulletin have gone on to become famous ornamentals, and a few have become infamous invasive species. To illustrate the latter category, I found this quote about Amur corktree in the June 14, 1911, issue of the Bulletin: \"Phellodendron sachalinense [now classified as P. amurense], which is a native of Saghalin [Sakhalin] and the northern island of Japan, has grown in the Arboretum into a tree about thirty feet high, with a tall, straight trunk, and widespreading branches for ming a shapely flat-topped head. The seed- The picturesque forms and thick, corky bark of the mature corktrees along lings springing up naturally near the Meadow Road often draw the attention of Arboretum visitors (Phellodendron old trees indicate that it is likely to amurense var. lavallei seen here). NANCy rOSE 6 Arnoldia 69\/2 ArCHIvES OF THE ArNOlD ArBOrETUM Weather talk one tHeMe that runs through the arboretum publications is a thorough discussion of weather and its effects on the collections. this seemed to be especially true for the Bulletin, which was aimed primarily at a local audience and often emphasized the immediacy of the growing season. take, for example, the october 16, 1929, entry by e. H. Wilson, \"the phenomenal drought which Massachusetts in common with other states has endured will long be remembered for it caused grave anxiety among all who garden. in late June the arboretum enjoyed one good rainfall but not another worth mentioning until october 2nd. For fully two months supplying water to suffering trees and shrubs was the principal work engaged upon. Fortunately, there was no great heat but at the height of the drought it looked as if a great many plant must die. thanks to the water stored from the heavy rains of spring the trees suffered but little and as autumn arrived a general freshening among all woody plants was noticeable. today it is difficult to realize that extreme drought has been experiThis photo by Donald Wyman shows tree damage at the enced. the power of resuscitation enjoyed by plants Arboretum from the hurricane of September 21, 1938. is, indeed, marvelous.\" Perhaps the most famous weather event to affect the arboretum was described by Wyman in the october 7, 1938, Bulletin: \"rain had been falling rather consistently for four days when on september twenty-first, over large areas in new england, the downpour assumed the proportions of a deluge. rivers in western Massachusetts were at flood stage, and everywhere the ground was soggy from excessive rain. by late afternoon the rain slackened and the wind increased to a gale. at 4:50 p.m. when the lights went out in the administration building staff members expected a \"blow\", but certainly did not anticipate the hurricane which caused frightful damage throughout new england. the arboretum lost approximately 1500 trees, and a recent newspaper estimate of the number of trees lost in Massachusetts--only one of the new england states touched by the storm--reached the appalling figure of 100,000,000. there is no way of checking such an estimate, but with definite information concerning the number of trees destroyed in a few boston suburbs, this figure seems possible... Hemlock Hill in the arboretum is one of the higher points between boston and the blue Hills. With wind velocities at times approximating 125 miles an hour it is understandable that great damage was done to the particular plantings on the southern or exposed side an the top of that hill. to the older friends of the arboretum, this damage will seem the most serious.\" (the tradition continues with an annual report in Arnoldia recapping the previous year's weather and its effects on arboretum collections.) science at the arboretum: seeing the Forest through the trees William (Ned) Friedman tHe arnold arboretuM is all about science, and has been since its founding in 1872 by Harvard university. the arboretum's mandate as stated in the original deed--to grow all of the trees and shrubs from anywhere on earth that could be grown here--was a long-term research proposal in itself, one that continues to this day. over the decades research in many fields has been conducted at the arboretum by our scholars as well as colleagues from institutions around the world. the opening of the Weld Hill research building early this year brings a new era of science to the arboretum. Weld Hill's state-of-the-art facilities include laboratories, greenhouses, and spectacular teaching equipment for undergraduates. Microscopes with lasers allow scientists to peer into the microscopic world of plants; molecular biology equipment allows us to unravel the dna that codes for the processes that make each plant and plant species unique and exquisitely responsive to its environment; and highly sophisticated banks of growth chambers permit botanists to study the effects of climate change on plants under controlled conditions. importantly, Weld Hill allows arboretum researchers formerly based at Harvard's camThe Weld Hill research facility at the Arnold Arboretum. bridge campus to expand their work at the arboretum. it also provides great new opportunities for students, scientists, and visiting scholars to conduct research using the living collections and the Weld Hill facilities. in essence, the arnold arboretum of Harvard university is poised to become a worldwide hub for the study of plant biodiversity. With over 15,000 curated living organisms, there are unlimited and unique opportunities to conduct botanical research at the arboretum. research has limited value until it is shared with others, of course. a vital part of the arboretum's mission over the years has been to translate the science of the arboretum to a wide audience. arboretum publications, especially Arnoldia and its predecessor, the Bulletin of Popular Information, have been important vehicles for disseminating information about the fascinating world of plant science to arboretum friends and colleagues around the world. as research grows at the arboretum we will continue to share it through Arnoldia as well as our much expanded website and education programs. William (Ned) Friedman is Director of the Arnold Arboretum and Arnold Professor of Organismic and Evolutionary Biology at Harvard University. JON HETMAN Bulletin of Popular Information 9 While these later publications were more complete and formal, the original Bulletin observations provide a more intimate connection to the seasonal cycles of the Arboretum. For me, the most remarkable thing about the Sargent and Wilson contributions is their timelessness. Having worked at the Arboretum for over thirty years, I can read their words and get the feeling that I can go right out onto the grounds and see the exact same scenes (or close to them) that they were describing. \"Bussey Hill, where the new and rarer plants from the Orient are quartered, is perhaps the most interesting place in the Arboretum at the moment [May 16, 1927]... From the overlook itself looking toward the south, the Hemlock Grove looms majestic; westward across the Oaks, over and beyond the steely gray, misty, cloud-like clump of American Beech, Spruce, Fir, and Pine stand conspicuous. Everywhere wholesome scented air, opening bud, blossom, and green grass--everything fresh and clean--the Arboretum in spring is rich in charm and beauty.\" When it comes to writing arti- ing this 102-year-old sand pear (Pyrus pyrifolia) in the open area just cles, no one was more prolific--in below the summit. both the Bulletin and Arnoldia-- cific purposes, based on his notes on the perthan Donald Wyman. He published hundreds formance of the collections over the years. In of pages of observations on the Arboretum's horticultural circles of the 1940s and 1950s, collections covering an amazing array of topics Wyman was the voice of authority in the including crabapples (his specialty), rhododennortheast when it came to recommending (or drons, lilacs, winter injury, hurricane damage, condemning) plants. trees with interesting bark, the order of bloom of Arboretum plants, seashore gardens, hedges, tHe Future vines, and a number of arcane topics such as In reading through Arboretum publications the use of spent hops for mulch (in Arnoldia from the past hundred years it's interesting to volume 7, Number 12, December 12, 1947). note how some things change and some stay More than any other of the Arboretum's horthe same. Many horticultural recommendaticulturists, he had no qualms about passing tions published in the Bulletin and Arnoldia judgment on the suitability of plants for spe- Bussey Hill still provides magnificent views and beautiful plants, includ- rICHArD SCHUlHOF 10 Arnoldia 69\/2 "},{"has_event_date":0,"type":"arnoldia","title":"Can Taxodium Be Improved?","article_sequence":2,"start_page":11,"end_page":20,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25520","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260b36d.jpg","volume":69,"issue_number":2,"year":2011,"series":null,"season":null,"authors":"Eguiluz-Piedra, Teobaldo; Yunlong, Yin; Zhou, Lijing; Creech, David","article_content":"Can Taxodium Be Improved? David Creech, Lijing Zhou, Yin Yunlong, and Teobaldo Eguiluz-Piedra COURTESy OF STEvEN J. BASKAUF, hTTp:\/\/BIOIMAgES.vANDERBIlT.EDU Growing in wetland conditions, this stand of baldcypress (Taxodium distichum) displays the distinctive knees (pneumatophores) characteristic of the species. D egradation of coastal forests and associated wetland habitats by excessive flooding and saltwater intrusion is a global problem, and may become even more so if predicted climate changes and consequent rises in sea level occur. In the United States, there's been great concern about the degradation of the entire Mississippi River Delta biotic system, much of which can be traced to manmade changes in the nature and flow of the Mississippi river. One example of this degradation is the loss of coastal forests south of New Orleans, which has left this city more vulnerable than ever to the impact of hurricanes. (Allen 1992; Earles 1975; Krauss et al. 1999) These circumstances make it increasingly important to identify, select, and even improve tree species that have some innate tolerances to flooding and salinity. Such trees will be valuable for restoring degraded coastal areas as well as for urban landscapes and other greening projects. For this reason, we are particularly interested in Taxodium distichum. Taxodium Taxa Of all native swamp forest tree species in the southern United States, Taxodium distichum (baldcypress) has long been recognized as being among the most tolerant to flooding and salinity. This long-lived and generally pest-free 12 Arnoldia 69\/2 Taxodium 13 All phOTOgRAphS By ThE AUThORS ExCEpT WhERE NOTED interfere with routine maintenance such as lawn mowing. While their exact function is unknown, knees may contribute substantially to wind throw resistance (Conner et al. 2002). Baldcypress in the western part of its range (central and western Texas) is generally more salt and alkalinity tolerant, and is less prone to produce knees than baldcypress from more eastern sources. East Texas genotypes of Taxodium planted in San Antonio, Texas, where soils are highly alkaline, often turn chlorotic and perform poorly. As with pond cypress and baldcypress, botanists and horticulturists speculate that baldcypress in central to western Texas are perhaps commingled with Montezuma cypress and represent transitional genetics (lickey and Walker 2002). This very large, old pondcypress grows at Shangri La Gardens, in Orange, Texas. While outside the normally accepted range for the species, this pondcypress appears to be spontaneous, not planted. Pondcypress occurs in the southern portion of the range of baldcypress and only on the southeastern coastal plain from North Carolina into louisiana. While southeast Texas is not normally included as part of the pondcypress natural range, an approximately 1,200-year-old pondcypress at Shangri la gardens, Orange, Texas, appears to broaden the range. pondcypress occurs in blackwater rivers, ponds, bayous, and swamps, usually without alluvial flood deposits. pondcypress is relatively easy to distinguish by its feathery foliage, which is ascendant, \" 14 Arnoldia 69\/2 Taxodium 15 distinct knees, is generally more tolerant of year, there are already apparent differences in salt and alkaline soils, and is less tolerant of foliage color, tree form, growth rate, and branchextended flooding. At Stephen F. Austin State ing characteristics. In Mexico, Montezuma University gardens in Nacogdoches, Texas, cypress is considered quite variable from one Montezuma cypress forces new growth early provenance to another and nursery conditions in the spring and continues to grow late into can greatly impact growth rate and form. The the fall. Observations of Montezuma cypress in viveros genfor nursery in Texcoco has grown USDA plant hardiness zone 8 (average annual Montezuma cypress for the last twenty years minimum temperature 10 to 20 16 Arnoldia 69\/2 ROBERT MAyER Taxodium 17 Planted at the Stephen F. Austin State University Gardens in March 2010 as small one-gallon-container plants, these specimens of \"merit\" clone T406 from China had a very fast growth rate. This photograph is from July 2011. to female flowers of `Nanjing Beauty' (then known as selection T302). Fifteen selections were made in 1995, with the main characteristics for selection being fast growth rate, dark green leaf color during the growing season, and red-orange leaf color in the autumn. Several of these clonal selections are now widely used in China. Additional Montezuma cypress Taxodium in China Taxodium varieties and hybrids play a very important role in the southeastern China coastal vegetation plan, particularly in the floodplains of the delta and associated bottomlands and estuaries of the yangtze River. The planting of coastal windbreak forests in this area was initiated in 2005. There are many reforestation projects under way on the mainland side of dikes that run along the sea, both north and south of the mouth of the yangtze. These projects have received massive provincial and federal financial support and millions of trees will be planted by midcentury. Taxodium hybrids have also found a place in many of the large parks being constructed in the major cities of southeastern China. as grand all Taxodium 19 IMPRovING Taxodium Several Taxodium germplasm collections exist in the southern United States but they remain relatively unexploited. In addition to the Taxodium collection at Stephen F. Austin State University gardens, Dr. Donald l. Rockwood, University of Florida, gainesville, Florida, manages a large planting of varied genotypes, many of which serve as seed sources for superior seedlings, with plantings that target tolerance of fly ash, salinity, or polluted soils. Dr. Ken W. Krauss, at the United States geological Survey, National Wetlands Research Center, lafayette, louisiana, is collecting seed from survivor trees in the Mississippi Delta that have been exposed to increasing inundation and salt surges (Krauss et al. 2000; Conner and Inabinette 2005). By cruising the massive \"ghost cypress forests\" (large stands of dead or declining baldcypress) of the southern delta, individual survivor trees can be found that perhaps have good resistance to subsidence and high salinity. Their progeny may offer promise for reforestation projects in marginal sites, and the opportunity for selecting superior clones is immense. Finally, Dr. Mike Arnold, Texas A & M University, College Station, Texas, has planted a large collection of baldcypress genotypes from across the South; the collection includes central and western Texas provenances, as well as a collection of Montezuma cypress from Mexico and southern Texas (McDonald et al. 2008) At the government nursery near Jinjiang (Jiangsu, China), I viewed over a million Taxodium cuttings in the one-acre field of propagation beds during a visit in September 2011. The nursery manager, Mr. Zho, employed a half-dozen ladies who used high-pressure hoses to hand mist the cuttings. Every day, for 8 to 12 weeks, each worker managed her own long run of propagation beds, dragging her hose and wand and waving a stream of mist over the crop. After each run, the ladies would rest and visit with each other, waiting until the beads of water on the cuttings had evaporated, the signal that it was time to repeat the process. When I asked why he used this strategy, Mr. Zho reflected that he had previously used automated boom misters on a timer, but he had found that the ladies knew better when the cuttings needed water--they had a feel for the crop--and rooting percentages were now very high. --David Creech 20 Arnoldia 69\/2 "},{"has_event_date":0,"type":"arnoldia","title":"The Writing of New Trees: Recent Introductions to Cultivation","article_sequence":3,"start_page":21,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25523","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260bb6f.jpg","volume":69,"issue_number":2,"year":2011,"series":null,"season":null,"authors":"Grimshaw, John","article_content":"the Writing of New Trees: Recent Introductions to Cultivation John Grimshaw Editor's note: New Trees: Recent Introductions to Cultivation is an extensive new reference manual published by the Royal Botanic Gardens, Kew, in 2009. In this essay, co-author John Grimshaw describes the process of putting such a detailed publication together. New Trees: Recent Introductions to Cultivation John Grimshaw and Ross Bayton Kew Publishing, Royal Botanic Gardens, Kew. 976 pp. 2009 ISBN 978 1 84246 1730 I n 2002 I was living and working in the Netherlands. One afternoon the phone rang and a richly English voice identified its owner as Giles Coode-Adams. Might I be interested in writing a book about recently introduced trees? Following early retirement from a career in finance, Giles had devoted his time to horticultural causes, first fund-raising for Kew and later becoming Treasurer and then President of the Royal Horticultural Society. Among his numerous commitments he was also Chairman of the International Dendrology Society's Scientific and Education Committee, reflecting his personal interest in trees and woody plants in general. While involved with Kew he had been the occasional recipient of young specimens grown from seed collected on expeditions, and indeed had taken part in a seed-collecting trip to Japan. The only problem was that, for so many of the recently introduced trees he was encountering, there was no useful literature to look them up in. His proposal to the committee that a book on such trees would be valuable was accepted and they started looking for authors. I am still not quite sure how they found me, as I was then by no means dendrological, but I had recently co-authored and edited a well-reviewed book on snowdrops (Galanthus) that may have had something to do with it. To cut a long story short, the committee decided I was the right person for the job. Generous sponsors provided funding and the task could begin. Setting ParameterS The assignment was to produce a book about all the tree species introduced to cultivation in recent decades, or those that had been in cultivation for longer but for which there was no good description. The standard reference in Britain is the venerated W. J. Bean's Trees and Shrubs Hardy in the British Isles, first published in 1914 and last updated in the 1970s to form the eighth edition. Its botanical descriptions are complemented by readable discursive text about the plant and its cultivation requirements, rather than the terse encyclopaedic format of, for example, Kr 22 Arnoldia 69\/2 ERICH G. VAllERy, USDA FOREST SERVICE, BUGWOOD.ORG New Trees 23 maritime climates of the north and the Mediterranean climate to the south. An obvious vegetational difference is the commercial cultivation of olives in the Mediterranean and this matches, with remarkable precision, the area of southern Europe experiencing USDA Zone 9 winter temperatures (average annual minimum temperature 20 to 30 24 Arnoldia 69\/2 New Trees 25 for recording details of trees, but one that really interested me was a large specimen of Larix sibirica whose label recorded that it had come from H. J. Elwes in 1900: I connected with this tree on several levels. Henry John Elwes (1846 26 Arnoldia 69\/2 New Trees 27 high enough and just far north enough for their plants to have a sporting chance of being hardy in milder areas. One of its special trees is what is known as Aesculus wangii, although the name is technically invalid, which produces nuts 10 centimeters (3.9 inches) across--they are a trophy in themselves. Upon germination, seedlings rocket up to over a meter (3.3 feet) within weeks. Its habit of coming into leaf early may be a problem, but the tree itself seems to be winter hardy in Britain, at least. Strange though it may seem, Australia was the next most prolific source for recent tree introductions. This apparent anomaly can be explained by the single genus Eucalyptus, although there are several Acacia and Callitris species plus odds and ends from other genera too. Although of negligible interest in much of the area covered, Eucalyptus has a devoted fan club in the milder parts (maritime Europe, the Pacific Northwest). These enthusiasts have been searching out populations in the coldest part of each species' range, a classic example of intelligent plant-hunting, unlike the usual grabbing of material from the first population found. Whether or not such sourcing has done them any good after a series of hard winters in both of these regions remains to be seen--many eucalypts have been devastated. As they grow so fast, however, it won't be long before we see another crop appearing. Eucalyptus nitens, the shining gum, holds the record for the fastestgrowing tree in Britain, achieving 20 meters (65.6 feet) in six years in Oxfordshire, making a splendid-looking tree in that time, but alas, it was killed last winter. Mexico has also been an important destination for plant explorers in recent decades. With its huge diversity of ecosystems this is hardly surprising. Conifers and oaks are the two most important groups of Mexican trees for temperate gardens, but there are hardy Cornus, Crataegus, Fagus, Platanus, and Tilia too. It is always astonishing to think of \"temperate\" genera occurring in the tropics, but there is a huge diversity of Magnolia in tropical America, and rainforest maples and oaks are diverse in southeast Asia. Half the diversity of Juglans (walnut) is found in the Neotropics! Most of these tropical species are too tender for tem- An impressively large nut from the tree known as Aesculus wangii (the name has not been validly published), native to Vietnam. perate horticulture, but this shouldn't prevent experimentation with species along the tropical fringes. Many Mexican evergreen oaks are proving to be remarkably tough, and even if they are defoliated by a severe winter will usually resprout next spring. Of all the new trees I studied for the book I was most impressed by Quercus rysophylla, an evergreen oak with big leaves that emerge red or bronze, and which forms a stately and handsome tree, hardy in western Europe, the southeastern United States, and the Pacific coast. Experts continue to discover new species of oaks in the United States and Mexico, usually isolated on obscure mountain ranges. One such is Q. acerifolia, from a few mountains in Arkansas, a handsome red oak with excellent autumn colors. liberated from its native habitat it is proving to be versatile, succeeding across the northern and eastern United States, including in the Arnold Arboretum. In general, trees from Chile and New Zealand are for specialist growers in mild moist climates, and one has to be bold to grow any African woodies: even species from the most temperate parts of South Africa and the high mountains of East Africa are very tender. But in the right conditions it is fun to try--and even JOHN GRIMSHAW 28 Arnoldia 69\/2 "},{"has_event_date":0,"type":"arnoldia","title":"A Year With the Apples of the Arnold Arboretum","article_sequence":4,"start_page":29,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25519","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260b328.jpg","volume":69,"issue_number":2,"year":2011,"series":null,"season":null,"authors":"Sax, Miles S.","article_content":"Miles Sax I have spent the past year as a horticultural apprentice working in the Malus (apple and crabapple) collection of the Arnold Arboretum, a collection that has long been recognized for its importance to the horticultural and scientific worlds. Because of the Arboretum's many introductions and broad distribution of both cultivars and previously undiscovered Malus species from wild origin, it has been hailed as the \"`Mother Arboretum' for flowering crabapples\" (Fiala 1994). Once celebrated in an annual event known as Crabapple Saturday, this collection remains popular with Arboretum visitors, especially during spring bloom and fall fruit display. The dynamic nature of arboreta, including ongoing change in the fields of taxonomy, nomenclature, and research technology, inevitably leads to the need for periodic large scale review of the plant collections. Although historically the Arboretum's Malus collection has been a high priority, in recent decades hor- An old poster touts the Arnold Arboretum's crabapple collection. PhOTOS By The AuThOr exCePT where nOTed Part of the Arnold Arboretum's Malus collection in spring bloom, including white-flowered Malus hupehensis (accession 324-55-B) at left. ArChIveS OF The ArnOld ArBOreTuM A Year With the Apples of the Arnold Arboretum 30 Arnoldia 69\/2 Apples of the Arnold Arboretum 31 were examined and data were taken on size, shape, and color of the fruit. In spring 2011 a similar review was done by recording the size, shape, and color of flowers along with other morphological characteristics. This information was compared to existing descriptive references. For cultivars, the late Father John Fiala's book Flowering Crabapples: The Genus Malus was the primary resource used in determining identity and exploring the ornamental history of the genus. Because there is no current monograph of Malus, for accessions of wild provenance we used the flora pertinent to the region in question, e.g., Flora of China. This initial review was done to verify the plant material and confirm its correct taxonomic identity, allowing us to update nomenclature and ultimately make sure our plants are correctly labeled. The petaloid stamens of this Malus sieboldii `Fuji' (accession 418-68-C) flower are an example of the importance of field observation in a curatorial review. These petal-like structures, viewed from afar, make the flowers appear to be double. This curious discovery was not noted in Fiala's previous descriptions of this selection. ArChIveS OF The ArnOld ArBOreTuM Plant detectives observing physical characteristics (flower color, leaf shape, etc.) is the most common way of identifying plants but it's not the only way. A trip to the Arboretum's curation department can give a researcher access to records detailing the origins or place of collection for many plants located on the grounds. Another resource is the cache of maps--many hand drawn--detailing the exact locations of plants come and gone. the herbarium in the Arboretum's hunnewell building and the main harvard university herbarium yield pressed specimens of wild plants that we enjoy as landscape ornamentals today. A trip to the Arboretum's library may uncover volumes of floras spanning the world, or historical horticultural periodicals detailing the latest plant introductions that were in vogue at the time. the library's extensive archives provide a wealth of original documentation such as ernest h. Wilson's journals and letters to Charles sprague sargent from his many far-flung plant collectors. to have these resources condensed in a single place allows for research and discovery to be made in a way that is unique to the Arnold Arboretum. The archives of the Arnold Arboretum hold a treasure trove of documents, including many examples of Ernest but sometimes even all these resources are not enough H. Wilson's correspondence. to make a definitive conclusion on a plant's true identity or origin. in recent years the ability to look at a plant's genetic makeup has proved to be a very powerful tool in plant systematics. to that end, the usdA Plant genetic resource unit in geneva, new York, has graciously offered to genotype a handful of the Arboretum's mystery Malus. When the results are returned we can look forward to some answers from this unique plant identification technology. 32 Arnoldia 69\/2 Apples of the Arnold Arboretum 33 Knowing what the collection actually comprises was the first step in being able to ask the question, \"what is the collection missing?\" There was no way to assess what additions would be required without first understanding all the options. The Arboretum's collections focus primarily on plants of documented origin, in particular wild provenance, so that became the framework for the assessment. we assembled a list of all known Malus species and their infraspecific taxa (e.g., subspecies, varieties) from around the world and compared it with our existing accessions, with the goal of identifying key areas where the collection lacked diversity. Armed with this information, the curation department now has a desiderata (wish list) of plants and we can request material from other botanical institutions and germplasm repositories. The list will also be useful when determining goals for future plant collecting expeditions. hortiCulturAl CAre of the ColleCtion horticultural care and maintenance of the Malus collection was a major part of my apprenticeship. One of my primary goals was to take a hard look at \"best practices\" for growing apples in order to develop an action plan that would reflect some of the new thinking on orchard cultural practices. The Arboretum collection grows primarily ornamental crabapples rather than eating apples, but many of the horticultural concerns are shared. eating apples have been cultivated for centuries in a variety of settings, so apple orchards provide an interesting model system for understanding how we manage human created plant ecosystems. Consumer interest in organic products has increased, as have demands that growers utilize better practices that are more environmentally friendly. what initially started with farmers is now spilling over into MIChAel dOSMAnn The author pruning in the Malus collection. 34 Arnoldia 69\/2 My favorite Malus nAnCy rOSe nAnCy rOSe the eCleCtiC mix of wild germplasm, hybrids, and early cultivars in the Arnold Arboretum's Malus collection gives inquisitive visitors a chance to see crabapples rarely found in the commercial trade. here are a few of my favorites: Malus `Mary Potter' FEATUrES: A medium height, wide-spreading tree with high disease resistance, offering abundant white flowers and small (0.4 inches [1 centimeter] diameter) red fruits. dESCrIPTIoN: This specimen is the original selection of the cultivar. Introduced by the Arnold Arboretum, this Karl Sax selection is considered by many to be his best Malus hybrid. Named in honor of C. S. Sargent's daughter, this hybrid is a result of cross between M. sargentii `rosea' x M. x atrosanguinea. ACCESSIoN NUMBEr: 181-52 B LoCATIoN: 51-SW orIgIN: Arnold Arboretum Malus kansuensis var. calva FEATUrES: rare in cultivation, its small stature and unique flowers and fruit make this an interesting apple in the collection. dESCrIPTIoN: Small, slow-growing tree; flowers are creamy white and fruits develop a caramel yellow color with a red cheek. The fruit is somewhat flattened on the top and bottom and has vertical ridges around it, giving it a pumpkin-like appearance. ACCESSIoN NUMBEr: 134-43 A LoCATIoN: 49-SE orIgIN: China Malus tschonoskii FEATUrES: Silver-white, tomentose undersides of leaves, attractive orange to red fall color, tall (40+ feet [12+ meters]) upright-pyramidal shape. dESCrIPTIoN: This accession is the Arnold Arboretum's oldest apple in the collection as well as one of the tallest. Collected by C. S. Sargent in 1892 during his expedition to Japan. The flowers and fruits of this specimen are insignificant, but the unique leaves and form look unlike any other apple. To the casual passerby it would be difficult to identify it as an apple tree at all. ACCESSIoN NUMBEr: 3678-A LoCATIoN: 17-SW orIgIN: Japan Malus hupehensis FEATUrES: The fruits are yellow with a red cheek and provide a nice contrast with the crimson to purple fall leaf color. dESCrIPTIoN: Wide-branching, vase-shaped tree. Leaves and copious fruit develop out of short branch spurs, giving a distinctive appearance. Leaves have reportedly been used as a tea substitute in parts of China. The species was introduced by the Arnold Arboretum and was first collected from China by E. H. Wilson in 1908. ACCESSIoN NUMBEr: 324-55 B LoCATIoN: 50-SW orIgIN: China Malus x robusta `Arnold-Canada' FEATUrES: A rare cultivar that is a towering giant of an apple tree. By far the tallest specimen in the collection. dESCrIPTIoN: Primary scaffolding branches alone are larger than the main trunks of many other Malus. The distinctive bark has an appearance somewhat similar to Prunus (cherry). This specimen features copious fruits that are orange-yellow with a bright red cheek. rarely found in other collections outside the Arnold Arboretum ACCESSIoN NUMBEr: 172-52 B LoCATIoN: 50-SE orIgIN: Hybrid 36 Arnoldia 69\/2 Apples of the Arnold Arboretum 37 This specimen of Malus spontanea (accession 10796-2-A, previously listed as Malus halliana var. spontanea) has an interesting shape and an interesting history. Perhaps because of its provenance or its status as an e. h. wilson-collected lineage it was preserved and is now growing perfectly well in its new orientation. Initially struck by its unusual form, I came to realize that this Malus had an interesting tale to tell. while conducting my curatorial review I was searching through our plant records in an effort to verify the identities of the Malus in our collection. My research brought me upon four living specimens of Malus halliana var. spontanea, all of which are the wilson lineage and one of which was the Malus in repose. looking at the provenance information I noticed that the original accession (10796-A) was wild-collected from Japan by wilson during his 1918 expedition. Although the taxonomy of Malus halliana is a bit unclear, what struck me as odd is that this species is reported as a native of China. My initial thought was that this was an accident in nomenclature and so I began to pursue the tree's true identity. Accession information stated that the plant was wild-collected by wilson, but without providing an exact location. Malus halliana has been cultivated as an orna- mental in Japan for generations, but since the Arboretum's specimens were supposedly from wild origin I realized something wasn't adding up and exact provenance information would have to be unearthed to get to the true identity of this specimen. weeks later, while conducting the conservation portion of my curatorial review, I was searching Malus on the Botanic Gardens Conservation International's (BGCI) website. To my surprise I saw that on the 1997 IuCn red list of Threatened Plants Malus spontanea (as a species, not a variety of M. halliana) was flagged as vulnerable. realizing that the Malus in question might be of conservation value, I decided I had to give another go at this mystery apple. I figured that if our records indicated that this plant was wild collected from Japan, somewhere buried in the archives there must be conclusive evidence of the true identity of this tree. After multiple searches in our herbarium, archives, and historic records, I eventually found the information I had been looking for in an article on new taxa by Alfred rehder in the Journal of the Arnold Arboretum (rehder 38 Arnoldia 69\/2 Apples of the Arnold Arboretum 39 Iketani offered to collect seed from the remaining wild populations and send them to both the uSdA and the Arnold Arboretum. Once this plant material clears the importation process we look forward to the infusion of these plants of high conservation value into the collection. ConClusion working as an Arboretum apprentice for the last year has been a fulfilling experience that has pushed me both intellectually and physically. The chance to engage with both the horticulture and curation departments led to many synergistic benefits. In 2010 I made 369 observations that resulted in data entries in BGBASe, and in 2011 I added another 560 observations, for a total of 929 observations on 479 individual plants. These data will be valuable for long-range curatorial planning as well as current horticultural maintenance, and may also be of benefit to fellow botanical institutions who hold Malus collections. My apprenticeship has been extended for another year so I will be able to continue my efforts to push this collection toward the highest levels of care and curatorial value. Bibliography Bunker, J. P. 2007. Not Far From the Tree. waterville, Maine: (self published). dosmann, M. S. 2009. Malus at the Arnold Arboretum: An Ongoing legacy. Arnoldia 67(2): 14"},{"has_event_date":0,"type":"arnoldia","title":"The Royal Azalea: Rhododendron schlippenbachii","article_sequence":5,"start_page":40,"end_page":40,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25522","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260b76b.jpg","volume":69,"issue_number":2,"year":2011,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"The Royal Azalea: Rhododendron schlippenbachii Nancy Rose R hododendron schlippenbachii is perhaps most noted for its lovely spring bloom, but this deciduous azalea is also a standout in the autumn garden when its leaves turn striking shades of yellow, orange, and red. In addition, royal azalea displays attractive summer foliage and a handsome winter silhouette, making it that object most desired by gardeners--a plant with all-season ornamental interest. Royal azalea has long been a favorite of mine, so I was tickled to find out that several illustrious Arboretum horticulturists have also written glowingly about this species. As Peter Del Tredici mentioned in the first article in this issue, many timeless bits of information and opinion can be gleaned by reading through old issues of the Bulletin of Popular Information and Arnoldia--here, Charles Sprague Sargent, first director of the Arboretum, describes royal azalea's native range and growth habit: R. Schlippenbachii is one of the commonest shrubs of Korea and often forms the dominant undergrowth in open woods. From Korea it crosses into northeastern Manchuria where it grows on the shores of Possiet Bay; it occurs, too, in two localities in northern Japan. Wilson found it extraordinarily abundant in Korea on the lower slopes of Chiri-san and on the Diamond Mountains, which were where he visited this region in June \"a wonderful sight with literally miles and miles of the purest pink from the millions of flowers of this Azalea.\" In Korea this Azalea on the wind-swept grass-covered cliffs of the coast grow[s] less than a foot high but flowers abundantly. In the forests of the interior it often grows to a height of fifteen feet and forms a tall and slender or a broad and shapely shrub. (Bulletin of Popular Information, May 5, 1921) Typically blooming in mid-May at the Arboretum, royal azalea is covered with large flowers in clear shades of pink, somewhat resembling a mass of pink butterflies resting on the branch tips. In the same Bulletin article quoted above, Sargent wrote, \"The pale pink fragrant flowers, which are about three inches in diameter and marked on one of the lobes of the corolla with red-brown spots, are perhaps more beautiful than those of any other Azalea, certainly of any Azalea which has proved hardy in the Arboretum.\" And Ernest H. Wilson wrote in the May 16, 1927, issue of the Bulletin, \"The blossoms on this lovely Korean Azalea are now open on the Bussey Hill. A sturdy bush of upright habit, bearing on naked twigs terminal clusters of large pale to pure pink blossoms. This is a very hardy and satisfactory Azalea.\" (Cold hardy through USDA Zone 5 [average annual minimum temperature -10 to -20"},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23426","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd1608128.jpg","title":"2011-69-2","volume":69,"issue_number":2,"year":2011,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Tree by Tree, Yard by Yard: Replanting Worcester's Trees","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25518","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260af6f.jpg","volume":69,"issue_number":1,"year":2011,"series":null,"season":null,"authors":"Freilicher, Mollie","article_content":"Tree by Tree, Yard by Yard: Replanting Worcester's Trees Mollie Freilicher T MASSAchUSETTS hiSTOricAL SOciETy cOLLEcTiONS he trees of Worcester, Massachusetts, have had a hard time recently, as Asian longhorned beetle (ALB) (Anoplophora glabripennis) has decimated public and private trees there and in surrounding communities. What may be less well known is that the invasion of ALB is just the most recent blow in what has been a difficult century for trees in the Worcester area. Some of the issues that the city has faced, such as canopy defoliation from gypsy moth (Lymantria dispar) and tree death from chestnut blight (Cryphonectria parasitica) and Dutch elm disease (Ophiostoma ulmi), were not unique to Worcester, affecting many communities across the eastern and central United States. Other threats have been more localized. Worcester, along with much of New England, suffered major tree loss in the 1938 Disasters have hit Worcester's trees before, including the devastating 1953 tornado. Photo of tornado damage by Alfred K. Schroeder, June 1953. Replanting Worcester's Trees MichAEL BOhNE, BUgWOOD.Org 3 PA DEPT. OF cONSErvATiON AND NATUrAL rESOUrcES, BUgWOOD.Org An adult male Asian longhorned beetle. ALB larvae tunnel into host trees, damaging essential conductive tissues. After pupating, adult beetles emerge from large, round exit holes, as seen on this maple trunk. hurricane, and a tornado in 1953 devastated parts of Worcester. More recently, the ice storm of 2008 damaged many trees, some beyond recovery, in the Worcester area. The ongoing onslaught has made it difficult for planting efforts to keep up with tree losses. Following the 1953 tornado and the already significant losses from Dutch elm disease, Worcester began growing maples and ramping up its planting efforts. Maples, especially Norway maple (Acer platanoides), known for its urban adaptability, became a mainstay of the planting program. Over the next six decades, maples came to comprise 80% of street trees, leaving many of the city's public trees vulnerable to a maple-specific insect or disease. Enter Asian longhorned beetle. This mapleloving insect was found in Worcester in August 2008 and has since been detected in four surrounding towns, spurring the creation of a regulated area in Worcester county that now measures 98 square miles. Like the hurricane and tornado, ALB quickly changed the land- scape of Worcester's northern neighborhoods. residents felt bereft of trees and looked to state and federal authorities to come forward with a solution. Even in areas without ALB, communities across the United States have been losing trees to development, to neglect over time, and to a lack of adequate replacement programs. Add up Worcester's experience with the hurricane, the tornado, the ice storm, and ALB, and the importance of replanting becomes clear. On a psychological level, replanting is also an important part of the healing process following the losses to ALB and the ice storm. Tree Benefits While urban residents have enjoyed shade and the aesthetic benefits of trees, in the last several decades researchers have studied the ecological, psychological, and social benefits of trees in urbanized areas. it is now known that trees are important for air quality, watershed health, carbon dioxide reduction, soil quality, noise reduction, property values, and psychological and 4 Arnoldia 69\/1 Replanting Worcester's Trees 5 experience. Some streets that had been lined with maples quickly became barren. residents felt the character of their neighborhoods had completely changed. images of empty streets were reminiscent of pictures following the tornado, when some streets were stripped of all trees. With the losses to ALB and the ice storm, the opportunity arose to reshape the urban forest--to improve diversity by planting a variety of non-host trees (trees that cannot support ALB), to move away from monocultures of maples, to strategically place trees along streets, in parks, and on private property to ensure that they have adequate growing space now and in the future, to educate residents about the value of trees and how to care for and maintain them, and to keep track of the new trees over time. The scale of such a replanting effort was larger than in any of the other ALB infested areas in the United States (parts of New york, illinois, and New Jersey). Planning the Plantings The lead federal agency in the Massachusetts ALB cooperative Eradication Program is the United States Department of Agriculture's Animal and Plant health inspection Service (APhiS) and the lead state agency is the Massachusetts Department of conservation and recreation (Dcr). replanting has been a component of all ALB programs in the United States and the United States Forest Service (USFS) is the lead agency for working with cooperators toward this end. Some cooperators in the Worcester area include municipalities and non-profit organizations that have stepped up MOLLiE FrEiLichEr Professional staff, volunteers, and property owners have all been involved in tree replanting efforts. Here a DCR team in Boylston plants trees. 6 Arnoldia 69\/1 MOLLiE FrEiLichEr Replanting Worcester's Trees 7 NANcy rOSE wrapped up in spring 2010. The goal of the USDA-funded planting was to mitigate the impact to the communities where host trees were removed because of ALB infestations. The USDA-funded planting specifically targeted property owners who lost trees to ALB in the 2-square mile core area where most removals occurred in 2009. For a property owner to be eligible for a tree with the USDA planting, the owner had to have lost a tree over six inches diameter at breast height (dbh) from a maintained area of the property. This put the focus on replacing landscape and specimen trees on properties in areas where natural regeneration could not be expected. Naturalized, unmaintained areas that could regenerate on their own were not included in this planting. Already two years on, property owners are seeing these areas come back to life. By spring 2009, the funding was in place from the USDA to plant approximately 800 trees in the areas first affected by tree removals. Properties that lost trees to ALB were identified from the USDA database and Dcr foresters mailed information to property owners about replanting. interested property owners responded and staff scheduled visits to select trees and locations. Additionally, staff went door to door to reach property owners who did not respond. As the USDA planting program was wrapping up, the next program was just getting started. The American recovery and reinvestment Act of 2009 (ArrA) provided $4.487 million in funding for the second planting program that got underway in spring 2010. The ArrA planting will continue into 2012. it targets all property owners in the regulated area regardless of whether they lost a tree to ALB. The only limit to the number of trees a property can have is the number of trees the property could support. in addition to increasing diversity and the number of trees on private property, the ArrA planting aims to restore public shade trees, to plant 15,000 trees on private property, to restore forest canopy and watershed functions affected by reduced canopy, and to create jobs. As of May 16, 2011, over 4,700 trees have been planted through the ArrA program. in addition to working with residents to site trees, Dcr foresters also conduct inspections of trees planted in previous seasons to ensure that trees Though popular ornamental trees, mountain ashes (Sorbus spp.) were not offered in replanting programs because of their suscebtibility to ALB. S. aucuparia `Michred' shown here. Host Genera for Asian Longhorned Beetle Acer Aesculus Albizia Betula Celtis Fraxinus Platanus Populus Salix Sorbus Ulmus Cercidiphyllum Koelreuteria Maple horse chestnut Mimosa Birch hackberry Ash Sycamore Poplar Willow Mountain ash Elm Katsura tree goldenrain tree 8 Arnoldia 69\/1 10 Arnoldia 69\/1 Trees selected for the replanting effort include (left to right) seviceberry (Amelanchier spp.), tuliptree (Liriodendron tulipifera), red oak (Quercus rubra), and linden (Tilia spp.). Table 1. Replanting List for Massachusetts Regulated Area. This represents current and past species that have been offered in the replanting program. LARGe SHADe TReeS Carpinus caroliniana American hornbeam Cladrastis kentukea (syn. lutea) yellowwood Fagus sylvatica European beech Ginkgo biloba* ginkgo Gleditsia triacanthos honeylocust Larix spp. Larch Liquidambar styraciflua Sweetgum Liriodendron tulipifera Tuliptree Metasequoia glyptostroboides Dawn redwood Nyssa sylvatica Blackgum Ostrya virginiana American hophornbeam Quercus alba White oak Quercus bicolor Swamp white oak Quercus coccinea Scarlet oak Quercus palustris Pin oak Quercus rubra red oak Tilia spp. Linden\/basswood ORnAMenTAL TReeS Amelanchier spp. Serviceberry Chionanthus virginicus Fringetree Cornus spp. Dogwood Malus spp. crabapple Prunus spp. cherry Syringa reticulata* Japanese tree lilac eVeRGReen TReeS Abies concolor White fir Picea pungens colorado spruce Pinus strobus Eastern white pine Thuja occidentalis* American arborvitae *No longer offered NANcy rOSE (ALL PhOTOS) 12 Arnoldia 69\/1 Replanting Worcester's Trees 13 MOLLiE FrEiLichEr Data on these recently planted street trees on Fairhaven Road in Worcester will be gathered for a central database that will help evaluate tree establishment and survival. software to explore the structure of and the environmental services provided by the newly planted trees. costs, and Strategic Planting. United States Department of Agriculture, Forest Service, Pacific Southwest research Station, general Technical report, PSW-gTr-202. herwitz, E. 2001.Trees at risk: reclaiming an Urban Forest. chandler Press, Worcester. http:\/\/www.beetlebusters.info\/ http:\/\/www.massnrc.org\/pests\/alb\/ i-Tree. n.d. http:\/\/www.itreetools.org\/ M. Freilicher, B.c. Kane, h.D.P. ryan, iii, and D.v. Bloniarz. 2008. Trees in Peril: responding to the Asian Longhorned Beetle. http:\/\/www. mass.gov\/dcr\/stewardship\/forestry\/urban\/docs\/ Worcester_report.pdf ric, J., P. de groot, B. gasman, M. Orr, J. Doyle, M.T. Smith, L. Dumouchel, T. Scarr, and J. Turgeon. 2006. Detecting Signs and Symptoms of Asian Longhorned Beetle injury: Training guide. Available online: www.glfc.forestry.ca\/vLF\/ invasives\/alhbdetecguide_e.pdf Worcester Tree initiative: http:\/\/www.treeworcester.org Mollie Freilicher is a Forester with the Massachusetts Department of conservation and recreation. What's next? As of May 24, 2011, the replanting program has planted nearly 6,000 trees and has found homes for over 9,000 trees for the spring and fall 2011 plantings. it will be some years before the streets of Worcester are lined with large trees again, but the diversity of trees that are being planted today will help buffer the city against future pests. Strategic placement of trees now can also help eliminate later conflicts with infrastructure such as power lines. With the many partners involved, and support at the state and federal level, Worcester and the rest of the regulated area is poised for an exciting recovery. References E. g. McPherson, E.g., J.r. Simpson, P.J. Peper, S. L. gardner, K.E. vargas, and Q. Xiao. 2007. Northeast community Tree guide: Benefits, "},{"has_event_date":0,"type":"arnoldia","title":"Geographic Information Systems for the Plant Sciences","article_sequence":2,"start_page":14,"end_page":22,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25516","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260ab26.jpg","volume":69,"issue_number":1,"year":2011,"series":null,"season":null,"authors":"Morgan, Brian J.","article_content":"Geographic Information Systems for the Plant Sciences Brian J. Morgan T he disciplines of the plant sciences and geography have been intertwined as far back as circa 300 BCE when the Greek scholar Theophrastus, frequently referred to as the \"Father of Botany,\" described the habitat and geographical distribution of plants in his first work on the subject titled Enquiry into Plants (Historia Plantarum). It wasn't until the sixteenth century and the establishment of the world's first botanical garden in Padua, Italy, that the leading icon of modern geography, the map, found its permanent place in the plant sciences by documenting the locations of woody plants in the garden for identification purposes. Today, location--the unifying theme of geography--has taken on an even more important role in the plant sciences where it is considered an essential attribute to record, and variable to consider, for the study of plants in fields ranging from agriculture to ecology. In the digital age that we live in, the cataloging of plants and the analysis of the influence that location plays on the growth and distribution of them is increasingly performed using geographic information systems (GIS). GIS is commonly defined as a system of personnel, computer hardware, software, and data for capturing, managing, analyzing, and displaying all forms of geographically referenced information. GIS merges the visual aspects of a map with the analytical power of a database, and allows plant scientists to view, question, understand, interpret, and visualize data in many ways that reveal relationships, patterns, and All ImAGES By ThE AuThor ExCEPT whErE noTEd Analysis of plant condition at the Arnold Arboretum reveals a cluster of plants in poor condition (indicated by red dots), in this case mostly eastern hemlocks (Tsuga canadensis) damaged by hemlock wooly adelgids. Geographic Information Systems 15 GIS is used to explore plant collections in public gardens. By linking maps with the collections database, details about accessions such as this white oak (Quercus alba) are readily available. trends in the form of maps, globes, reports, and charts. In our rapidly changing world, GIS gives scientists the power to quickly understand and formulate solutions to the problems presented by our most complex issues such as population growth, resource consumption, and climate change. ComPonentS of a GIS Personnel and Equipment The foundation of a powerful GIS is built with the personnel required to develop and manage the system. GIS managers and analysts usually have a strong background in the principles of cartography and database management systems, and a number of graduate and certification programs have appeared in the last decade to support this education. Fortunately, recent advances in simple desktop GIS software like Esri ArcGIS Explorer for the visualization and analysis of any geographic data, and in webbased solutions like Google maps, have made the use of GIS technology accessible to all. with expert personnel in place, the next item needed for a geographic information system is the computer software used to capture, manage, analyze, and display spatial data. desktop GIS software packages like Esri ArcGIS desktop or Quantum GIS are most commonly used to perform the majority of GIS functions, but serverbased systems like map Server and Esri ArcGIS Server are increasingly being employed to share data, maps, and even analysis capabilities with users through their web browsers without the need for training or special software. These server systems even allow for the collection and use of GIS data and maps on mobile devices like smartphones and tablets that contain location sensors such as a global positioning system (GPS) receiver. The computer hardware used for a desktop GIS is largely dependent on the requirements of the software selected, the amount and intricacy of the data to be analyzed, and the complexity of the analyses to be performed. while a standard desktop or laptop computer can be used for most systems, workstation-class desktop computers with fast processors, ample memory and storage, and high-performance graphics are most common. Server-based systems that allow multiple users to work with GIS data stored in a 16 Arnoldia 69\/1 Geographic Information Systems 17 as a conservation organization for biodiversity assessment, or a botanical garden for plant collection curation, it is common to design or employ an existing data model. A data model can be thought of as a database design or template that carefully considers how real-world features are represented as geometry in feature classes, the attributes appropriate for each feature, and any known relationships that exist between individual features or entire feature classes. In the plant sciences, user community designed data models for Esri ArcGIS exist Collecting GPS data at the UC Davis Arboretum. for biodiversity assessment, forestry, and public gardens. These models can be downloaded for free and allow scientists open Source vs. Commercial to get started with their GIS projects quickly, Software without the need to design their own modthere is much debate within the GIS develels. data models like the one developed by oper and user community regarding the the Alliance for Public Gardens GIS (ArcGIS Public Garden data model) additionally prochoice of commercial software versus open vide standardization across multiple organizasource software. Commercial products like tions, thus simplifying the exchange of critical those offered by esri and mapInfo are quite biodiversity data. expensive to purchase and typically require whether using an existing data model or annual maintenance fees for support and designing a new GIS from the beginning, one of the first things to consider is the availability upgrades, but offer well-designed user interof the data required for the project. A common faces and sophisticated analysis tools. Constarting point is to collect as much existing data versely, open source solutions such as those as is readily available through internet data that are part of the open Source Geospatial repositories like the united States Geological foundation (oSGeo) are free to use, but are Survey Earth Explorer or local government spatial data clearinghouses. one of the primary more difficult to operate and get support for. sources of this data is from a technique called Developers and scientists within academia remote sensing which is formally defined as generally tend to favor the use of open the collection of information about an object source software for GIS applications and without making contact with it. [Ed. note: See research, while private companies and govnext article for more on the use of remote sensing.] As it relates to spatial data, remote sensing ernments usually use commercial products. usually refers to data captured from aircraft or Ultimately, the choice is a tradeoff between spacecraft, and typically comes in the form of cost and ease of use, but the same functionaerial photography, multi-spectral images that ality is available from either option. measure non-visible forms of electromagnetic radiation, or even lIdAr height data that is CourTESy oF mIA InGolIA, uC dAvIS ArBorETum 18 Arnoldia 69\/1 Geographic Information Systems 19 Global greenness (vegetation) can be evaluated with the Normalized Difference Vegetation Index (NDVI) from remote sensing data gathered by satellites. sciences. The next section describes how it is being employed in real-world projects for both research and management. aPPlICatIonS of GIS our ever increasing need for land and resources combined with the threat of climate change has pushed the assessment of biodiversity to the top of the list of plant science research priorities. The scientific literature is rich with articles on the subject, ranging from studies of parks and reserves to the entire planet. GIS is often cited as the primary tool used to perform many of these studies, which frequently employ species occurrence data from informatics sites like the Global Biodiversity Information Facility (GBIF) and remotely sensed data from satellites like landsat 7 Tm to determine the relative species richness of a particular area. In one study of African vascular plant diversity, the investigators performed a multivariate analysis to determine the relationship between the number of species in well-known areas and the associated environmental conditions like topography, temperature, precipitation, and evapotranspiration. This relationship was then used to interpolate the species richness in lesser-known areas to produce a vascular plant diversity map for the entire continent (mutke et al. 2001). once the biodiversity of an area has been assessed, the spatial data generated from the assessment can be used to help prioritize which parcels of land should be designated as conservation areas. Since the study of land and the process of delineating boundaries are inherently spatial in nature, GIS is cited as the overwhelming choice of tool for the task. In addition to biodiversity data similar to that produced in the previous study, conservation planning activities usually include topography, precipitation, soil, geology, and land use data. In a forest conservation study in malaysia the investigators used a decision making approach that assigns weighted values to possible alternatives in an effort to prioritize areas for conservation. This study considered species and ecosystem diversity, the soil and water conservation functions of plants, and potential threats to the forest, and through a process called map algebra, hot spots for conservation were determined and used to delineate potential new protection areas (Phua and minowa 2005). GIS can also be an invaluable tool when planning a collecting expedition. Traditional approaches to expedition planning have favored areas that were considered interesting or easily accessible, and tended to focus on species that were easily studied. GIS allows for the unbiased Geographic Information Systems 21 GIS data can be represented as points, lines, polygons, and rasters. Here spot elevations are shown as points, contours are shown as lines, ponds are shown as polygons, and an elevation surface is shown as a raster in part of the Arnold Arboretum. sampling of an area that not only maximizes heterogeneity, but can also assure that the largest gaps in the record of biodiversity are filled with the least amount of effort and resources. GIS-based expedition planning typically utilizes existing biodiversity data along with topography, geology, vegetation, temperature, and precipitation data to determine areas that have the greatest potential to provide the maximum amount of new information and specimens. In a survey gap analysis study in Guyana the investigators used museum and herbaria specimen data to locate geographical gaps in the existing data in an effort to determine candidate survey sites for each taxonomic group of interest. These candidate sites were then compared with weighted abiotic variables to determine a final set of collecting sites that had the greatest chance of producing new information and specimens for each taxonomic group (Funk et al. 2005). once plant specimens have been collected and cultivated in a botanical garden or arboretum, GIS is commonly used to curate the collection and to help make management decisions throughout the entire lifetime of the plant. GIS is commonly used for creating collection maps and planting plans, identifying problems and threats, planning mitigation, and performing research. In addition to data about the living collection, a typical garden GIS employs data on topography, soils, hydrology, land use, facilities, transportation and more. In a tree conservation study at the uC davis (university of California, davis) Arboretum the investigators used data on the location, species, size, and condition of each specimen in conjunction with data on site characteristics and conflicting urban infrastructure to determine a condition rating and a hazard rating for each tree in the collection. These ratings were then used to identify areas of concern and to produce prioritized mitiga- Facing page: Thematic layers in the ArcGIS Public Garden Data Model allow comparisons and correlations of databases from soils and topography to plants and animals. 22 Arnoldia 69\/1 "},{"has_event_date":0,"type":"arnoldia","title":"Remote Sensing as a Botanic Garden Tool","article_sequence":3,"start_page":23,"end_page":32,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25517","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260ab6b.jpg","volume":69,"issue_number":1,"year":2011,"series":null,"season":null,"authors":"Griffith, Patrick; Witcher, Ericka","article_content":"Remote Sensing as a Botanic Garden Tool Ericka Witcher and Patrick Griffith MONTGOMeRy BOTANICAL CeNTeR R emote sensing is a tool already in use for plant exploration, ecology, forestry, habitat restoration, and other related fields. It also has great potential in botanic gardens for botany, horticultural science, and management purposes. At Montgomery Botanical Center, located in Coral Gables, Florida, we were able to improve our assessment of the property with the addition of new software that provided the capability for deeper evaluation of the collections and natural resources using remote sensing imagery and data. By adding LIDAR (Light Detection And Ranging) imagery to maps and employing techniques normally used at larger regional scales, new information was discovered about the garden and its collections. Garden maps serve multiple purposes. Their primary use is as a location catalog--what a garden has and where it is. People who use the garden, whether staff or visitors, will want to know where certain features are at some point. The information displayed in this kind of map can reflect the vastly different purposes of, say, a researcher examining different subspecies of Coccothrinax miraguama (miraguama palm), an irrigation techni- A 2008 aerial photo of Montgomery Botanical Center property in south Florida. cian repairing a break, or a visitor through the addition of a spatial or geographic looking for the restroom, but all three of their component--where things are in relation to garden maps would need to show what things something else. Spatial relationships in a botanare and where they are located. On the other ical garden, for example, can examine how close hand, maps can also be used for more dynamic vulnerable plants are to open spaces or high-use purposes in the garden. New areas of horticulvisitor areas, how tree canopies change over tural and scientific interest can be illuminated 24 Arnoldia 69\/1 Remote Sensing 25 eRICkA WITCHeR Vegetative and geologic characteristics, like canopy and elevation changes seen here along the Palm Walk, are quantifiable with LIDAR-integrated maps. origins or sensitive nature of many plants in our collections, we must work to create and maintain an environment that provides for their individual needs for life and growth. To that end, we are continually looking for new ways to assess the garden property and analyze both its biological and geological resources. Legacy Imagery For several years we had utilized aerial photographs to examine tree canopy and other features at MBC that were difficult to thoroughly evaluate from ground level. Orthophotographs (planimetrically-corrected aerial photographs) and uncorrected aerial photos are frequently used in many different industries, including botanic gardens, for many disparate purposes, and are readily available through a variety of sources (e.g., the USGS website http:\/\/www. usgs.gov\/pubprod\/, or state or county websites). These photos provided a good general sense of how areas were developing, but we experienced a fair amount of difficulty integrating them with our AutoCAD (a computer-aided-design software program)-based maps, so their utility was somewhat limited. We wanted a way to view the photos and the maps at the same time as well as use other types of imagery, then be able to perform spatial analysis. New Systems Add Capability A software grant for botanical gardens and zoological parks provided an all-in-one solution. Two MBC staff members had prior experience with the software, and with the help of an additional intern, by late 2009 we had completely converted the old maps and their CAD layers to a GIS (geographic information system). The local coordinate system was replaced with geographic latitude and longitude so the con- 26 Arnoldia 69\/1 Remote Sensing 27 By eRICkA WITCHeR FOR MONTGOMeRy BOTANICAL CeNTeR height and density were examined in the LIDAR images and transects were distributed and performed accordingly. The invasive plant was not found to be as pervasive as feared, and as a result, eradication efforts were scaled down proportionally (edelman and Griffith 2010). Using LIDAR imagery to better visualize the dense plant growth beforehand gave us a more complete picture prior to entering the area, saving time and effort. For another project we adapted a conventional forestry analysis using first-return LIDAR images to appraise height and breadth information (Sumerling 2010) to establish potential candidates for national or state champion tree status. (Champion trees are the largest known individuals of a species based on measurements of height, trunk circumference, and canopy spread.) This was done by simply overlaying the plant layer over the image and visually identifying the tallest canopies. The plant curators also applied their infield knowledge of the various species' usual growth habits Bare-Earth LIDAR image of MBC property showing ground level surface geology; to propose more individulower (darker) areas west of the escarpment are important for planting, as they als for assessment, the height are more likely to contain sand and silt, in contrast to the surrounding alkaline and spread of which were also limestone bedrock, or the clay marl to the east. checked in the LIDAR map. A first-return LIDAR image also offered a At writing, 27 trees had been awarded state lot of utility for other vegetation-assessment champion status by the Florida Division of projects. First-return images illuminate all the Forestry, and 2 trees received national chamtopmost surfaces of the study area; in this case, pion status from the conservation organization canopy height and coverage. In one project, an American Forests. undeveloped section of the property filled with Future Development both an invasive exotic plant, Schinus terebinWith the successful completion of these projthifolius (Brazilian peppertree), and protected ects, we have become more familiar with the mangrove trees needed a thorough evaluauses of both aerial photography and LIDAR for tion so we could determine the most efficient horticultural purposes, and subsequently the course of action for managing the land. Canopy 28 Arnoldia 69\/1 By eRICkA WITCHeR FOR MONTGOMeRy BOTANICAL CeNTeR Remote Sensing 29 30 Arnoldia 69\/1 By eRICkA WITCHeR FOR MONTGOMeRy BOTANICAL CeNTeR eRICkA WITCHeR Remote Sensing 31 This Florida champion tree, Pterygota alata (Buddha coconut), has endured dozens of hurricanes, and at 89 feet tall is one of the tallest trees on our property. We first identified it as a candidate through examination of LIDAR imagery. eRICkA WITCHeR potential applications for this kind of data. We are also exploring additional applications currently in use by other landscape-level industries that have a good deal of potential for use in botanic gardens (Perroy et al. 2010, Sumerling 2010). For example, we are now working on creating contours from a first-return LIDAR image that will provide new information about the canopy coverage and biomass density in the garden. This type of three-dimensional data can amplify current knowledge about shade structure, wind protection, and plant growth and expansion within the property and create an operational image of the \"vegetative topography.\" Coupled with 15 years of database records tracking the growth and reproductive activity over time of our plants, we anticipate new insights to spur in-depth research. LIDAR also lends itself to three-dimensional modeling and creating fly-throughs, leading to comprehensive visual aids for online garden \"explorers\" and researchers, as well as garden managers looking to gain new perspectives on their collections and resources. Meanwhile, maximizing survival rates of invaluable scientific plant collections with analysis of current collections and records, both spatial and temporal, is an ongoing objective. By employing imagery in our GIS and adapting some of the more basic and conventional uses of LIDAR for regional landscapes to the localized, relatively small-scale botanical garden, we have been able to save many hours of laborious fieldwork and gain a nuanced understanding of the property and plants under our care. Acknowledgements: We thank Brian Witcher and Judd Patterson of the National Park Service SFCN program for their GIS assistance and consultation, MBC GIS Intern Jonas Cinquini and MBC Intern Sara edelman for their project assistance, the Stanley Smith Horticultural Trust for all GPS equipment funding, and the eSRI Botanical Garden\/ Zoological Park Grant program for providing all GIS software. LIDAR imagery provides clues to planting conditions at MBC for staff biologist, Chad Husby, looking for future plant sites near the all 32 Arnoldia 69\/1 "},{"has_event_date":0,"type":"arnoldia","title":"Book Review: Weeds: In Defense of Nature's Most Unloved Plants","article_sequence":4,"start_page":33,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25515","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260a76d.jpg","volume":69,"issue_number":1,"year":2011,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Book Review: Weeds: In Defense of Nature's Most Unloved Plants Peter Del Tredici Weeds: In Defense of Nature's Most Unloved Plants Richard Mabey. Ecco, an imprint of HarperCollins Publishers. 324 pp. 2010 (United States publication 2011) ISBN 978-0-06-206545-2 I n his new book Weeds: In Defense of Nature's Most Unloved Plants, Richard Mabey presents a refreshingly non-judgmental look at some of the most vilified plants on earth. While acknowledging the problems that some of these notorious plants can cause for both gardeners and ecosystems, he also presents their not insubstantial positive contributions in terms of recolonizing derelict land in cities, restoring war-ravaged landscapes in Europe, and, over the millennia, providing abundant food and medicine for people. In short, the author takes a balanced approach to the subject of weeds and he puts the focus where it belongs--on their intimate association with human culture going back to the dawn of agriculture itself. As Mabey presents it, the subject of weeds is nothing less than a microcosm of human culture, an observation that he reinforces with numerous quotations from famous writers including Shakespeare, Ruskin, and Thoreau, and, of course, from the Bible. Not stopping here, he also provides a lengthy discussion of the significance of weeds in visual arts, as exemplified by a discourse on the significance of Albrecht D 34 Arnoldia 69\/1 Book Review 35 aggressive invasive species such as giant hogweed (Heracleum mantegazzianum) and kudzu (Pueraria montana). It should also be noted that the book is up-to-date in its discussion of the modern, scientific data on weeds, discussing in detail how the increased use of herbicides over the past fifty years has influenced weeds' evolution, and how genetically modified (GM) crops are interacting with weeds to make them hardier and more difficult to eradicate. In short, Mabey masterfully weaves the disparate fibers that constitute the cultural and natural history of weeds into a colorful tapestry of a book that few nature writers can match. Weeds: In Defense of Nature's Most Unloved Plants is not without a few flaws however, one of which (for American readers) is its exclusive use of the British common names of plants throughout the text. There is a glossary at the end which provides the Latin equivalent to the common name, but the fact that many of the plants discussed in the book have different common names in North America than they do in England leaves the inquisitive American reader who doesn't know the Latin names of plants with little choice but to turn to the internet or reference books to figure out identities. In addition, the book is overwhelmingly focused on weeds that dominate the landscapes of the British Isles and on British writing on the subject, making the book somewhat less relevant to North American audiences than it perhaps needs to be. Certainly the history and behavior of North American weeds is discussed in the book, particularly the subject of their early introduction from Europe, but their treatment is minimal compared to the space devoted to weeds in Britain. There's also a surprising absence of any mention of the extensive pioneering German literature on the subject of urban ecology, particularly that done by Herbert Sukopp and his colleagues in post-war Berlin. Despite the British focus of Weeds: In Defense of Nature's Most Unloved Plants, I found it a fascinating read--which is no small accomplishment given the fact that I have a large library of well-studied weed books at home. Mabey is an engaging writer with long-standing, highly personal interest in weeds that shines through on Dandelion (Taraxacum officinale). every page. He deserves kudos for his masterful integration of the scientific and cultural aspects of weed ecology and his fluid, often poetic, use of language. Here he describes watching weeds grow at an active construction site: \"When I look at their comings and goings, as hectic as the movements of the bulldozers, I grope for metaphors to understand their meaning. I think of ants, but they're too organized, too determinedly earth-changing, like the excavating machinery itself. Then it occurs to me that they are like a kind of immune system, organisms which move in to repair damaged tissue, in this case earth stripped of its previous vegetation.\" While this book has something for everyone, I suspect that its greatest appeal will not be to down-in-the-dirt gardeners but to those of the armchair persuasion who like their weeds with a touch of literature, humor, and taste. Peter Del Tredici is a Senior Research Scientist at the Arnold Arboretum. NANCy RoSE "},{"has_event_date":0,"type":"arnoldia","title":"A Venerable Hybrid Oak: Quercus x sargentii","article_sequence":5,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25514","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260a728.jpg","volume":69,"issue_number":1,"year":2011,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"A Venerable Hybrid Oak: Quercus x sargentii Michael S. Dosmann S cores of plant taxa--species, infraspecific variants, and hybrids--commemorate Charles Sprague Sargent with their epithets. They range from the cherry palm of the Caribbean, Pseudophoenix sargentii, to the vase-shaped Sargent cherry of East Asia, Prunus sargentii. In 1915, yet another plant was given the Sargent moniker when Arboretum taxonomist Alfred Rehder recognized the Arboretum director by providing a name for the hybrid between the English oak, Quercus robur, and the American chestnut oak, Quercus montana (formerly known as Q. prinus). While hybrids between these two members of the white oak subgenus (Lepidobalanus) had been known since the 1830s, this was the first time the taxon was recognized officially with its own name, Quercus x sargentii, the Sargent oak. From Q. robur, the hybrid attains a certain nobility and majesty, not to mention a girthy trunk, broadly spreading canopy, and distinctive auriculate (earlobe-shaped) leaf bases. From Q. montana come the crenately toothed leaves, smaller-stalked acorns, and, with age, coarsely furrowed bark. The Sargent oaks that grow in the Arboretum's living collections can all be traced to the initial lot of acorns collected from a magnificent tree at Holm Lea, Sargent's estate in Brookline, Massachusetts. The seeds arrived at the Arboretum on October 6, 1877. They germinated and yielded multiple seedlings that were planted in the permanent collections and cataloged under accession number 5883. Currently, three plants (A, B, and C) remain in the collection, each looking exceptional for being over 130 years old. Perhaps the most spectacular is 5883-A, a majestic specimen located near the junction of Bussey Hill Road and Beech Path, at the base of the Forsythia and Syringa collections. With a current height of 84 feet (25.6 meters) and DBH (diameter at breast height) of 55.7 inches (141.5 centimeters), this tree commands attention. Visitors strolling down Beech Path often pause in awe to admire the tree's massive limbs and rounded crown. Recent landscape renovations to this area, known as State Lab Slope, will not only maintain the health and vitality of this specimen and the surrounding plantings, but also improve visual access. I should note that its siblings (plants B and C) may be slightly smaller, but are also notable and worth a visit. Both are located further along Beech Path, near the edge of the Fraxinus collection. Q. x sargentii is extremely rare in cultivation, and our understanding of it is essentially limited to the specimens grown in our collection as well as those of a few other botanical gardens and arboreta. Certainly, our three trees are exceptional and have stood the test of time, but it would be premature to say much more without further study. I am particularly interested in this hybrid's potential use as a tree tolerant of the vagaries of the managed landscape, especially in urban areas where soils are prone to drought and other limitations. As Q. montana is an upland species typically found growing in dry and rocky habitats, one could hope that the Sargent oak is similarly tough. Oaks are difficult to propagate clonally, and attempts over the years to clone the Arboretum's trees have been in vain. However, because Q. robur is a species that can sometimes be rooted from cuttings, Manager of Horticulture Steve Schneider and I are conducting several experiments to see if ease of propagation from this parent was passed along to the hybrid. If that is the case, it opens up a great deal of potential for additional study and, perhaps, the Sargent oak's use as a street tree near you. Michael S. Dosmann is Curator of Living Collections at the Arnold Arboretum For additional information on this hybrid and its interesting history, see: Hay, I. 1980. Outstanding plants of the Arnold Arboretum: Quercus x sargentii. Arnoldia 40(4): 194 "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23425","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160bb6e.jpg","title":"2011-69-1","volume":69,"issue_number":1,"year":2011,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Tapping the Underappreciated Plant Diversity of the Eastern United States","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25512","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070896b.jpg","volume":68,"issue_number":4,"year":2011,"series":null,"season":null,"authors":"Lewandowski, Rick J.","article_content":"tapping the underappreciated Plant Diversity of the Eastern united states Rick J. Lewandowski T All pHoToS By RICk J. leWANDoWSkI exCepT WHeRe NoTeD he romance and intrigue of plant discovery and acquisition continues to entice plant explorers, most often to remote and exotic places far away from the United States. Though early explorers and botanists (including the Bartrams, the Michauxs, Nuttall, Torrey, Gray, and Harper) described the vast richness of eastern North America's flora, its range of diversity and adaptability continue to be underappreciated and understudied to this day. In efforts to more fully document and explore its potential, Mt. Cuba Center, near Wilmington, Delaware, is among a handful of public gardens currently active and engaged in exploring and promoting this rich flora. Plant ExPloration with PurPosE The forests of eastern North America are replete with a remarkable array of plant communities, habitats, and plant species. Through the vision and resources left by our founders, Mt. Cuba Center has dedicated significant energy The Dogwood Path forms an intimate canopy adjacent to Mt. Cuba Center's meadow. It showcases many plants in sophisticated layers, from tall trees to low-growing wildflowers. Plant Exploration 3 Gorge rhododendron (Rhododendron minus) is one of the native species being documented, collected, and evaluated. The wild population seen here on Flag Mountain in Alabama is at home in the hot and humid southern end of its range on shady, dry, acidic slopes. to exploring habitats throughout the eastern United States from pennsylvania and Delaware through the Carolinas, Georgia, and Alabama. Throughout these states we have discovered that there is an enormous reserve of genetic diversity worthy of greater study and appreciation. plant exploration is an essential component of Mt. Cuba Center's commitment to study and assess the adaptability of native plants for horticultural use. In addition to enriching the gardens with documented, wild-collected, seed-grown plants from a range of provenances, detailed field data gathered from this effort has provided us with a greater understanding of habitats, distribution, and plant associations. This documentation has potentially far-reaching implications for horticulture, landscape design, and conservation. During the past 11 years, Mt. Cuba Center staff members have conducted nearly 80 field expeditions in the eastern and southeastern United States in 11 states. over 1,150 documented collections have been made, representing 619 taxa of herbaceous and woody plants. Regular collaboration with numerous partners--including other public gardens, universities, state and federal agencies, industry, conservation organizations, and private individuals--has afforded us the opportunity to observe and sample plant diversity in a wide range of habitats. In addition to broad-based sampling of herbaceous and woody taxa, Mt. Cuba Center's field work in recent years has also focused on sampling specific taxa in order to obtain broader genetic diversity, obtain taxa from the edges of their ranges or from disjunct populations, and assess potential variation of selected plant species for wider landscape use. Some of the highest priority woody plant taxa for targeted sampling currently include: Fothergilla gardenii, Fothergilla major, Halesia carolina, Halesia diptera, the Copeland Family legacy MT. CUBA CeNTeR is the 589-acre former estate of Mr. and Mrs. lammot du pont Copeland. over a 65-year period, beginning in 1937, gardening became a consuming passion for the Copelands. During the formative years of their estate and garden, the Copelands engaged a number of designers including Thomas Sears, Marian Coffin, and Seth kelsey to assist in creating a series of formal and informal landscapes surrounding their home. These provided structure and a unique identity to their ever-expanding gardening interests. By the late 1970s Mr. and Mrs. Copeland had begun to refine their gardening interests, focusing more and more on native plants of the eastern United States. With passion, vision, and hands-on garden development, the Copelands and their staff, including their first director, Dr. Richard lighty, created some of the mid-Atlantic region's most attractive and diverse native plant gardens. In 2002, a year after Mrs. Copeland's death at age 94 (predeceased by Mr. Copeland in 1983), Mt. Cuba Center officially became a private non-profit organization, with a mission to display, study, and promote the broader use of the flora of the eastern United States, with particular emphasis on the piedmont physiographic region. Today, a decade later, well documented and botanically diverse gardens, horticultural research and introduction programs, as well as extensive education and public tour programs are beginning to fulfill the Copelands' hopes for their beloved estate. The Copelands' Colonial Revival home was completed in 1937 and today serves as the hub of administrative and education activities at Mt. Cuba Center. In the foreground, a dense stand of native prickly-pear cactus (Opuntia humifusa) spills over a rocky ledge. Plant Exploration 5 Halesia tetraptera, Illicium floridanum, Kalmia latifolia, Leucothoe axillaris, Leucothoe fontanesiana, Rhododendron catawbiense, Rhododendron colemanii, Rhododendron minus, Rhododendron prunifolium, Stewartia malacodendron, Stewartia ovata, Viburnum acerifolium, and several deciduous Rhododendron species. somE ExamPlEs oF initiativEs CurrEntly unDErway: Sampling Stewartia Diversity Silky camellia (Stewartia malacodendron) and mountain camellia (Stewartia ovata) are attractive native North American deciduous shrubs or small trees that have been underappreciated for their horticultural value and as biological indicators of stable, botanically rich habitats. Both species have highly attractive, non-fragrant, white flowers that open daily over a one to three week period from mid May to late June, depending upon the region. The distribution of these two species is often discontinuous, Silky camellia (Stewartia malacodendron) flowering in mid June at Mt. Cuba with highly variable popula- Center is a show-stopper. tion sizes. Silky camellia is primarily found in the coastal plain and piedmont species overlaps, with plants of both species from Virginia to Florida and west as far as eastgrowing in close proximity to each other. ern Texas. Mountain camellia is naturally disIn 1999, Mt. Cuba Center began targeted tributed in the piedmont and mountains from field work to document populations and collect Virginia to Georgia and Alabama, reaching its seeds of both species from across the breadth of western limit in southern kentucky and Tentheir ranges. More than 120 documented seed nessee; a few disjunct populations can be found collections of these two species from 7 states in the coastal plain of Virginia and North Carohave since been made. lina. While there are pockets of large populaThe Stewartia Working Group (SWG) was tions, most populations are small and isolated, formed in 2007 as a collaboration to develop ex which is likely to have resulted in genetic isositu repositories for the extensive documented lation. North-central Alabama is one (possibly seed collections already made and to study, the only) place where the range of these two long-term, the variation in these two eastern 6 Arnoldia 68\/4 These four images show variation in the filament color of Stewartia ovata flowers on separate plants. North American species. The SWG currently includes the Birmingham Botanical Garden, Alabama; Mt. Cuba Center, Delaware; polly Hill Arboretum, Massachusetts; Smithgall Arboretum, Georgia; yew Dell Gardens, kentucky; and Heritage Seedlings nursery, oregon. Much of the success of the SWG is due to the knowledge, guidance, and assistance of stewartia authority Jack Johnston. He has systematically identified countless stewartia populations and guided the group to them in order to observe and sample both silky camellia and mountain camellia. As a result, a significant portion of the range of these two species has already been sampled. Much remains to be learned, though, about the variation, habitats, distribution, propagation, production, and adaptability in cultivation of both species. even so, the SWG is making significant progress in representing a broad range of native stewartia diversity in cultivation. provenance-based collecting such as that with Stewartia malacodendron and Stewartia ovata offers an important window into the variation of species and potential preservation of genetic variation. It also affords emergent opportunities to encourage research as well as selection of superior forms. Some other important provenance-based collections of interest include Leucothoe axillaris (6 collections), Leucothoe fontanesiana (14 collections), Rhododendron colemanii (8 collections), and Rhododendron prunifolium (7 collections) to mention just a few. In many cases these taxa are represented in cultivation by few or no known wilddocumented populations. Documented collections in our public gardens are crucial for Plant Exploration 7 Stewartia ovata is frequently found in rich mesic woodlands of the upper Piedmont and mountains of the southeastern United States where it can become a small tree. The specimen seen here is growing in Georgia's Warwoman Wildlife Management Area. expanding our appreciation of the variation in these species, properly identifying cultivated forms and hybrids of species, and providing documented, known-source material for horticulture and science. on thE EDgE Unfortunately, much of our pre-conceived bias about plant adaptability is based upon limited experiences with plants from their core ranges. Assumptions about adaptability become rules regarding how plants perform in the landscape, but are not always correct. In eastern North America, the range of many species is frequently broader than we know and is not fully represented in cultivation. Success and failure with a number of native plants in the garden at Mt. Cuba Center has informed our opinions about the need to more thoroughly explore the distribution of native eastern North American herbaceous and woody plant species. As a result, in the past several years field work has focused on locating, documenting, sampling, and growing a number of species from the edges of their ranges. Rhododendron minus Gorge rhododendron (Rhododendron minus) is found in the mountains of North Carolina, South Carolina, Georgia, and Tennessee, growing on well-drained slopes and outcrops. However, there are also many discontinuous populations of this species found into southern Alabama and Georgia. The location and environmental conditions of these southern populations places them under significantly greater heat and drought stress for extended periods throughout the growing season than mountain populations. 8 Arnoldia 68\/4 Alabama populations of Rhododendron minus frequently produce attractive pink flower trusses. While gorge rhododendron and its close relative, Carolina rhododendron (Rhododendron carolinianum), are attractive broadleaved evergreen shrubs that have been exploited for breeding and selection for many decades, their summer adaptability to landscape stress in the mid-Atlantic has been suspect. over the past several years, Mt. Cuba Center has made 26 collections of Rhododendron minus from throughout the southern end of its range to observe garden adaptability and variation; this work continues. It is interesting to note that nearly all the populations of gorge rhododendron from the southern end of its range develop attractive pale to dark pink flower trusses that bloom later than mountain populations despite their southern nativity. Through the guidance of rhododendron expert Ron Miller, we have also obtained and grown seeds from isolated populations of whiteflowering forms found in Alabama and Georgia. It is our hope that long-term observation and assessment of these plants may yield opportu- nities for wider introduction of these southern genotypes into cultivation as well as breeding and selection work. Additionally, the inclusion of these plants in our living collection provides opportunities for continued taxonomic study of this interesting group of rhododendrons. Rhododendron catawbiense Catawba rhododendron (Rhododendron catawbiense) generally ranges from the mountains and upper piedmont of the Carolinas and Tennessee northeast into Virginia and West Virginia above elevations of 3,000 feet (914 meters). Because of hot and humid summers (especially the warm night temperatures) in the mid-Atlantic region, this rhododendron struggles to survive in the typical suburban landscape. Again, with the knowledge and assistance of noted azalea and rhododendron expert, Ron Miller, we have been able to document and collect seeds from 12 populations along the extreme southern edge of the range of this rhododendron. In remote and difficult-to-access Creating a haven for herbaceous Plants MT. CUBA CeNTeR'S garden and living collection is an integrated matrix of plants intended to delight and inspire guests. It is our goal to provide guests with an understanding of how beautiful gardens are created and maintained using environmentally appropriate landscape management practices. Through this approach the garden and living collection provides opportunities for sophisticated layers of herbaceous plants that play an important role in the structure of the garden, integrating with and complementing the woody plant layers throughout the growing season. our gardens are well-known for an extraordinary variety of herbaceous plant species and cultivars that contribute character to the woodland garden setting. of Mrs. Copeland's favorite wildflowers, trillium (Trillium spp.) was the queen. To this day, trilliums continue to be among the most coveted of all wildflowers grown in our garden. over the past 20 years, we have developed expertise in the propagation and production of trillium species from seed. In 2001, Mt. Cuba Center was recognized for significant plant expertise with trillium propagation, production, and ex situ preservation by receiving North American plant Collections Consortium (NApCC) member status as an official holder for the genus Trillium. At present, the collection includes 84 taxa represented by more than 470 accessions. To broaden the genetic diversity of trilliums available in cultivation and to support ex situ preservation, we have continued to target trillium in our plant exploration activities. over the past 11 years, we have made 110 collections of wild-documented trillium species, varieties, and unique forms. These include: Trillium catesbaei, T. cernuum, T. cuneatum, T. decumbens, T. discolor, T. erectum, T. flexipes, T. lancifolium, T. luteum, T. nivale, T. rugellii, T. stamineum, T. underwoodii, and T. vaseyi. A number of hybrids and unique forms of several trillium species have also been collected. While trillium is an important focus, much work at Mt. Cuba Center continues on a very diverse assemblage of native herbaceous plants in order to contribute to the richness and diversity of plants worthy of wider use by the gardening public. Trillium decumbens Trillium simile Trillium discolor 10 Arnoldia 68\/4 BoTH IMAGeS By FReDeRICk R. SpICeR, JR., BIRMINGHAM BoTANICAl GARDeN altitude montane habitats farther north. evaluation of seedlings from these populations is in the early stages, but as with gorge rhododendron, observation and assessment may provide opportunities for broader use of this species in cultivation along the mid-Atlantic seaboard or for incorporation into breeding schemes. Kalmia latifolia Mt. Cuba Center has had a long-standing love affair with mountain laurel (Kalmia latifolia) because natural populations occur on the property and, more importantly, because the Copelands used this attractive broadleaved evergreen frequently in their garden. Mountain laurel is common throughout the eastern and northeastern United States all the way to Maine. Interestingly, it is also distributed into southern Alabama, the Florida panhandle, and eastern louisiana. While much selection and breeding work has been done with mountain laurel in the northeastern United States, there has been a limited emphasis on documenting, collecting, growing, and assessing the adaptability of mountain laurel from the extreme southern end of its range. Unlike populations of Kalmia latifolia in our area that Catawba rhododendron (Rhododendron catawbiense) flowers heavily even in dense grow in dry, shady upland woodshade in Alabama. However, its flowers are pale lavender pink rather than the lands, populations of mountain deeper violet purple of northern populations. laurel in the Deep South are frequently found in sandy, well-drained, shady sites along rivers and ravines in northeastern riverine habitats, sometimes well within the Alabama, Catawba rhododendron grows on flood zone of streams. Recently, 20 collections sandy benches at elevations ranging from 600 of mountain laurel from Alabama, Florida, and to 1,200 feet (183 to 366 meters). The climate louisiana were made to assess the long-term of the Alabama Catawba rhododendron habitats stress tolerance of these provenances compared is much warmer and more humid than higher Plant Exploration 11 In the southern end of its range Kalmia latifolia is frequently found in riverine habitats where flooding is common. Flowers of a Kalmia latifolia specimen growing in Coosa County, Alabama. 12 Arnoldia 68\/4 Mountain laurel was among the Copelands' favorite shrubs, serving as a year round evergreen that explodes into flower in early summer. to widely cultivated forms. This work is in its infancy but represents an important long-term opportunity to assess adaptability, disease and pest resistance, and ornamental value. assEssing aDaPtability Most gardeners in the mid-Atlantic region are unfamiliar with Florida anise (Illicium floridanum). For those who are, they usually consider it to be, at best, marginally hardy in our area. Despite this fact, Mrs. Copeland grew this lovely broadleaved evergreen shrub in her garden for nearly two decades. Its supple evergreen foliage, modestly formal upright habit, adaptability to shade and drought, as well as its attractive burgundy red flowers in early spring have made Florida anise a modern favorite of staff and visitors to this day. Florida anise is common in bottomland forests, along lakes and streams, and on the edges Dark red flowers of Florida anise (Illicium floridanum) stand out in the spring landscape. of wetland communities throughout much of southern Mississippi and southern to centralnorthern Alabama. Despite its common name, Plant Exploration 13 A flowering dogwood (Cornus florida) blooms in the spring mist along the Woods Path at Mt. Cuba Center. Florida anise is found only in the panhandle of Florida. In order to more systematically assess the garden adaptability and hardiness of Illicium floridanum, we began sampling populations on the northern edge of its range in Alabama several years ago. Through this work, 11 documented collections have been made, including samples from disjunct populations at the northern edge of the species' range in Alabama. Through long-term evaluation and distribution, we hope to broaden the potential for using Florida anise in the mid-Atlantic and surrounding regions. what's nExt? The flora of the eastern United States still has much to offer. The plants mentioned here are just a few of the many that deserve broader long-term study. Through Mt. Cuba Center's long-term commitment to observing, documenting, and sampling the flora of the eastern United States, we hope that a broader segment of this flora will be appreciated and used by the gardening public. Rick J. lewandowski is Director of Mt. Cuba Center near Wilmington, Delaware. "},{"has_event_date":0,"type":"arnoldia","title":"Leaf-out Dates Highlight a Changing Climate","article_sequence":2,"start_page":14,"end_page":22,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25510","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070856d.jpg","volume":68,"issue_number":4,"year":2011,"series":null,"season":null,"authors":"Primack, Richard B.; Polgar, Caroline","article_content":"Leaf-out Dates Highlight a Changing Climate Caroline Polgar and Richard Primack T RichaRd PRiMack and caRoline PolgaR he arrival of spring is heralded each year by striking displays of flowers on trees and shrubs. Perhaps less conspicuous than the blooming of flowers, the emergence of new leaves on woody plants marks the onset of the growing season and controls a host of ecosystem functions. While to the untrained eye it may seem as though the leaves come out at the same time each year in one big burst, there are actually relatively consistent differences among species in leaf-out dates, as well as large differences from year to year in the timing of leaf out. The study of the timing of leaf out (and other natural annual phenomena) is known as phenology. Much of what we know about the physiology of leaf and bud development, and the mechanisms behind various leaf-out strategies, comes Clockwise from top left: Birches tend to leaf out early in the spring, while hickories tend to leaf out in late spring. Red oaks and beeches tend to leaf out in the middle of spring. Leaf-out Dates 15 caRoline PolgaR as red maple (Acer rubrum), white oak (Quercus alba), and red oak (Quercus rubra) (o'keefe 2010). The Japanese Meteorological agency has been recording leaf out, flowering times, autumn leaf color, and other phenological data of individual marked plants in phenological gardens at over 100 weather stations since 1953 (ibanez et al. 2010). ginkgo (Ginkgo biloba), also known as maidenhair tree, is one of the species being monitored. The international Phenological gardens (iPg) project, a network of botanical gardens across europe, has been collecting similar data on leaf-out dates Young red maple (Acer rubrum) leaves begin to expand in the spring. of individual plants since from research done in the past by tree physi1951 (Menzel 2000). other leaf-out datasets go ologists and foresters who were interested in back even further, including one that henry the connections to tree growth and timber hardavid Thoreau compiled in the mid-nineteenth vests. over the past few years, however, the century on plants in concord, Massachusetts. range of people interested in leaf-out phenolFindings from these studies indicate that there ogy has grown, as have the methods employed can be large year-to-year variability in the timto study it, largely as a result of the relevance ing of leaf out, depending on the weather, and of this phenomenon to global climate change. that there tends to be relative consistency in new technology, including satellite data, is the order of leaf out of species from year to year now being used to monitor leaf-out timing over (lechowicz 1984). wider areas than was possible in the past. Forest obtaining annual observations of leaf-out and ecosystem ecologists are connecting these dates can be quite time and labor intensive, leaf-out date observations to larger issues of often limiting studies to a small area around a global climate change, with implications for field station or a small number of species. To carbon cycles, the availability of fresh water, measure leaf out on a larger scale, remote sensand wood production. ing has emerged as a valuable new tool that can monitor an entire plant community or ecoMonitoring leaf out system consisting of many different kinds of The recent resurgence of interest in the phenolplants. Remote sensing studies typically use ogy of woody plants has led to leaf-out monidata obtained by sensors on orbiting satellites, toring projects around the world. For example, such as the advanced Very high Resolution over the past twenty years dr. John o'keefe and Radiometer (aVhRR) and the Moderate-resoother ecologists at the harvard Forest in central lution imaging Spectroradiometer (ModiS), or Massachusetts have been recording the dates of equipment on landsat satellites. Satellite sysleaf emergence of individual trees and shrubs tems vary in their spatial resolution, frequency each spring, including such common species of coverage, and types of data gathered. Scien- 16 Arnoldia 68\/4 FiSheR and MuSTaRd 2007 A 110 DateofAverageGreenLeafOnset(P ) 165 Km 0 25 50 100 B Km 0 50 100 200 The average onset of leaf out in (A) southern New England from Landsat (1984? 2002) and (B) the northeastern United States using MODIS (2000?2005). These images demonstrate that later phenology occurs at higher elevations, such as the Adirondacks and White Mountains; at higher latitudes; and in coastal areas that experience moderating ocean effects, such as Cape Cod and the Islands. The Boston and New York metropolitan areas leaf out earlier because of higher temperatures associated with the urban heat island effect; earlier leaf out is also seen in warm river valleys. Colors indicate the date on which half of the tree canopy has leafed out (from day 110 [April 20] to day 165 [June 15]), with earlier onset shown by blue and later onset by orange and red. (image from fisher and mustard 2007) tists use data transmitted from these satellites to calculate the changes in the amount of green vegetation (greenness) there is in a certain area over a growing season. analysis of graphs of greenness over time can be used to quantify important dates in the growing season, such as date of first leaf out in spring, the date at which half of the leaf cover has developed, and canopy senescence in autumn. Several recent research papers have shown that regional leaf-out data from satellites accurately match ground observations. This is particularly important because there is concern that different topographic features, such as mountains, fields, cities, and lakes, might create errors in the detection of green-up dates. in one study from Rhode island, researchers using landsat data were able to incorporate landscape features into their analysis and detect a delay in leaf out at the base of hills due to cold air drainage, a delay in coastal areas due to the cooling effects of the ocean, and a one-week delay in leaf out for deciduous forests in rural areas compared to those in the nearby urban area of Providence (Fisher et al. 2006). While such changes in leaf-out dates are already known from specific ground observations, the ability to detect such effects using remote sensing greatly extends our ability to map leaf out over large areas. another interesting remote sensing approach for monitoring leaf out uses phenocams, which refers to cameras placed in fixed locations that are used to record images of the leaf canopy at regular intervals, such as every hour or once a day, throughout Leaf-out Dates 17 couRTeSy oF John o'keeFe and The haRVaRd FoReST A sequence of two photos taken at the same spot by a phenocam at the Harvard Forest showing leaf out over a one-week period. The picture on the top left was taken on April 30, 2009, the one on the top right was taken on May 7, 2009. The photograph on the bottom was taken on April 29, 2009 and shows a view over the canopy of the Harvard Forest with the phenocam below. the growing season. dr. andrew Richardson and others have set up phenocams in the canopy at harvard Forest and 11 other forests in the northern continental united States. These images can be analyzed using computer programs to determine the seasonal trajectory of budburst, green-up, and senescence. networks of these phenocams can fill in the spatial and temporal gaps between plant monitoring by human observers and regional remote sensing images. Seven of these sites in the united States also have towers that monitor the exchange of carbon dioxide (co2) and water between the atmosphere and the forest. This combination of data from webcams, satellites, and gas sensors is providing crucial information on the relationship between phenology and ecosystem processes, especially carbon uptake (Richardson et al. 2009a). Leaf out and climate change climate change is already affecting many ecological processes, and leaf out is no exception (ibanez et al. 2010; Menzel 2000; Richardson et al. 2006). By analyzing long-term data on leafout dates, much can be learned about how the onset of spring has changed over time as temperatures have increased. From data collected at the iPg, researchers determined that trees in northern europe have advanced their leaf out by an average of one week over the past fifty years (Menzel 2000). in Japan, woody plants such as forsythia (Forsythia viridissima var. koreana), ginkgo, mulberry (Morus bombycis), and various cherry species (Prunus spp.) leafed out an additional 2 to 7 days earlier for each 1?c increase in temperature between 1953 and 2005. at a few sites, however, ginkgo trees were actually leafing out later than they did in the past, contrary to expectations (ibanez et al. 2010). at the hubbard Brook experimental Forest in new hampshire, the leaf out onset of three native species--american beech (Fagus grandifolia), sugar maple (Acer saccharum), and yellow birch (Betula alleghaniensis)-- has advanced an average of 5 to 10 days over the past five decades (Richardson et al. 2006). historical datasets, such as those recorded by henry david Thoreau and aldo leopold, can 18 Arnoldia 68\/4 also be used for these types of studies by comparing their records to contemporary observations from the same place, even if there is a lack of data between the two time periods (Bradley et al. 1999; Miller-Rushing and Primack 2008). We know that leaf out has become earlier in many areas in recent years, largely because of warmer temperatures, but what about the future? Will the advance in spring's onset continue for all species? To answer these questions, it is necessary to both be familiar with the physiology behind leaf out, and to build on what we already know about the response of leaf out to temperature. Variation in leaf-out times Trees and shrubs vary widely in leaf-out times, both among and within species. For instance, individuals will leaf out earlier in a warm, sunny location, such as a south-facing hill, than individuals of the same species located in a cold, shady location. Similarly, all individuals of a given species will leaf out later during a cold spring than in a warm spring. Sometimes when a tree is growing on the edge of field, the exposed sunny side will leaf out earlier than the shady side. These differences aside, there is a fairly consistent pattern in the leaf-out timing of trees, shrubs, and vines from year to year. RoBeRT MayeR Buttonbush (Cephalanthus occidentalis) is one of the latest native shrubs to leaf out in the spring. It is seen here blooming in midsummer. in eastern Massachusetts, species leaf out over a 4 to 6 week period. among the first plants to leaf out in the spring are such introduced ornamental shrubs as common lilac (Syringa vulgaris), honeysuckles (Lonicera spp.), and Japanese barberry (Berberis thunbergii), and nonnative fruit trees such as apple (Malus spp.). of native species, meadowsweet (Spiraea alba var. latifolia), quaking aspen (Populus tremuloides), black cherry (Prunus serotina), and grey birch (Betula populifolia) are among the first species to leaf out. consistently among the last species to leaf out are white ash (Fraxinus americana), white oak, and black tupelo (Nyssa sylvatica), with poison sumac (Toxicodendron vernix) and buttonbush (Cephalanthus occidentalis) often being the last of all. The pattern of leaf out is fairly consistent across the temperate zone of europe and north america. certain groups of plants tend to leaf out early (birches, willows, alders, many poplars and aspens) and others late (hickories, walnuts, and ashes). So why do some species leaf out so early and other species leaf out so late? Since the function of leaves is to carry out photosynthesis and provide sugars for the tree, in general it should benefit a tree to leaf out as early as possible to get the longest growing season. a tree species that leafs out in early april has four additional weeks to photosynthesize compared to a tree species that leafs out in early to mid May. however, the early-leafing tree faces the danger of a late frost that will kill its leaves and damage its vessel elements, the chief water conducting tissue. This trade-off between the advantages of early growth and of late growth provides a good explanation of why certain species leaf out when they do. The stem anatomy supports this explanation, with early species tending to have smaller vessel elements that are less prone to frost damage than the larger vessel elements of later species (lechowicz 1984; Miller-Rushing and Primack 2008). also important is the evolutionary history of a plant group: if it orginated in a warmer climate, it may not have fullyadapted mechanisms for dealing with extreme cold and therefore may have different factors regulating leaf out than a plant group originating in a colder climate. The vulnerability of trees and other plants to frost damage was recently demonstrated when Leaf-out Dates 19 ioWa STaTe uniVeRSiTy A late frost killed the new foliage on this oak tree. two weeks of abnormally warm weather in March 2007 triggered early leaf out all across eastern and central north america. a return of freezing weather from april 5 to 9 killed young leaves and flowers, and caused the die back of tree canopies across the region (gu et al. 2008). This frost damage was an example of the type of episodes of mismatches between plants and climate that may become increasingly common as climate change continues. What triggers leaf out? leaf out is predominantly controlled by temperature, with plants generally leafing out earlier in warmer conditions, but warm temperature is the not only factor. in fact, for many species it is a combination of warm and cold temperatures along with day length that dictates when the leaves will emerge from the bud. Most temperate species, including sugar maple and quaking aspen, have a chilling requirement, meaning that a certain number of cold (gener- ally a minimum of 0 to 10?c [32 to 50?F]) days in winter are required before the buds are able to break dormancy. The exact number of chilling units required depends both on species and on the weather of the preceding growing season (hunter and lechowicz 1992; Perry 1971). once this requirement has been fulfilled, a certain number of warm days above a certain temperature threshold are then needed for leaf development to begin and buds to open. This pattern is seen in both deciduous and evergreen species. in addition, some species also have a photoperiod requirement, meaning that they will only leaf out once daylength reaches a certain number of hours in the spring. in particular, longlived trees of mature forests, such as american beech, some oak species, and hackberry (Celtis occidentalis), often rely on a combination of photoperiod and temperature cues to break dormancy. For these species, budbreak only occurs after specific photoperiod and temperature requirements have been met. This holds even 20 Arnoldia 68\/4 when individuals from these species are planted in subtropical climates with exceptionally high temperatures (korner and Basler 2010). in contrast, many opportunistic species that are found early in forest succession, such as birches, hazelnuts (Corylus spp.), and poplars, do not have a photoperiod requirement to break winter dormancy. This somewhat risky strategy allows trees to respond more quickly to episodes of warm temperature in the early spring, but also creates more susceptibility to late frosts. yet a third group of species, which includes mostly ornamental plants from warmer climates, has a leaf-out strategy linked to spring temperature with minimal chilling requirements and no photoperiod requirement. The common lilac is a local example, and is one of the first plants to leaf out each spring. Learning from the past, predicting the future using information about past phenological responses to temperature and future climate scenarios, scientists can develop models to predict future phenological changes both at the species and ecosystem levels. one modeling study found that the advance in leaf-out time for most species and places is likely to continue in coming decades as the climate continues to warm (Morin et al. 2009). Many temperate tree species will show large advances in leaf out at higher latitudes, including the northern united States and canada. delays in leaf out, or abnormal leaf-out events, could occur at the southern end of species ranges in the southern united States for some species including black ash (Fraxinus nigra) and sugar maple if those species fail to meet their winter chilling requirement. Species with photoperiod requirements are also unlikely to continue to show linear advancements in leaf-out dates with increasing temperatures since photoperiod will not change. Because there are a host of complicated factors involved in leaf-out phenology, it is hard to predict whether leaf out will continue to advance linearly with changes in temperature at the whole forest level. The possibility of shifts in species composition resulting from climate change, as some species expand their range and others contract theirs, adds yet another layer of uncertainty to the prediction of leaf-out dates. if certain early successional species with minimal photoperiod and chilling requirements continue to leaf earlier in the spring, they may increase their abundance and distribution to become the dominant species, and shift the leaf-out time of the whole forest. The unmet chilling and photoperiod requirements of other species may significantly slow the advance of leaf out at the whole forest level. These two scenarios have consequences for many ecosystem processes, including the uptake of carbon dioxide, tree growth, forest temperature, and water movement. The earlier leaf-out times of many escaped ornamental shrubs, such as Japanese barberry and several honeysuckle species, may help to explain why these species are increasing so greatly in abundance in our forests. Their earlier leaf-out times may give them a competitive advantage over native species with more restrictive requirements for leaf out (Willis et al. 2010). Ecological interactions The onset of spring affects not only plants and ecosystem processes, but also organisms that depend on those plants. leaf-out timing determines the availability of food and shelter for many species, particularly insects. This timing is especially important for species that have gone through a long winter with little available food, or for bird species completing an energy-demanding migration north. From his close observations of nature in concord, henry david Thoreau was aware of the ecological importance of the emergence of leaves in the spring, writing in 1854: \"To-day the air is full of birds; they attend the opening of the buds. The trees begin to leaf, and the leaf-like wings of birds are in the air. The buds start, then the insects, and then the birds.\" Thoreau was aware of order of events based on what he had experienced in previous years and took for granted that the same pattern would persist, even with the large inter-annual variation in weather. in the twenty-first century, we can no longer take for granted that this order of natural events will continue each year. While plants are responsive to changes in temperature, other organisms that interact with plants Leaf-out Dates 21 RichaRd PRiMack Clockwise from top left: Photos of the Old North Bridge in Concord, Massachusetts, showing the development of the leaf canopy in the spring of 2010 (April 15, April 20, May 3, May 13). The meadow in the foreground is flooded in the first two photos, and dried out in the second two photos. in an ecosystem may not be quite so quick to respond. For instance, while certain species of birds arrive earlier in warmer years, other birds do not change their arrival dates, and some species are even arriving later (Miller-Rushing et al. 2008). insects are involved also: if certain kinds of insects feed only on the young leaves of a particular plant species that are present for a limited time in the spring, those insect species may decline in abundance if they emerge too early or late in the spring relative to their food resource. Birds that depend on those insects for food may similarly decline in abundance. Much more work is needed to understand how climate change and rising carbon dioxide concentrations are affecting ecosystem processes. if trees are leafing out earlier in the spring and dropping their leaves later in the autumn, they are likely increasing the net amount of carbon being sequestered in biomass. This possibility is supported by the work done by andrew Richardson and others at the harvard Forest and the howland Forest in Maine, showing that the earlier onset of spring in new england results in an increase of carbon sequestered in deciduous forest and somewhat less in coniferous forest (Richardson et al. 2009b). if this is occurring over a large area and over many years, the longer growing season could allow temperate forests to withdraw more carbon dioxide from the atmosphere. The longer growing season and warmer temperatures could also mean that 22 Arnoldia 68\/4 trees are losing more water vapor to the atmosphere during the process of photosynthesis; in consequence, forest ecosystems could possibly release less water to streams and aquifers, with major implications for drinking water supplies, flood control, and ecology of aquatic organisms. Conclusion Throughout the world, forests are being altered in many ways by the rising temperatures associated with global climate change, and the earlier leaf-out dates of trees and shrubs is one such example. earlier leaf-out dates are expected to continue in coming decades across much of north america. over a longer period of time, many tree species will likely be extirpated on a local scale and shift their ranges in response to the changing climate. Because there is a wide variation among species in leaf-out times, changes in the species composition of a forest will also mean changes in leaf-out dates at the level of the whole forest. disentangling the separate effects of changing species composition and changing climate is one of the great challenges of detecting leaf-out trends using remote sensing. Botanical gardens such as the arnold arboretum can contribute to these efforts by quantifying the differences among species in leaf-out dates for trees, shrubs, and vines all growing at one location, which for many species is outside of their native range. Such information can then aid in calibrating leaf-out dates over a large area using remote sensing. References Bradley, n. l., a. c. leopold, J. Ross, and W. huffaker. 1999. Phenological changes reflect climate change in Wisconsin. Proceedings of the National Academy of Sciences of the United States of America 96: 9701?9704. Fisher, J. i., J. F. Mustard, and M. a. Vadeboncoeur. 2006. green leaf phenology at landsat resolution: Scaling from the field to the satellite. Remote Sensing of Environment 100: 265?279. Fisher, J.i. and J.F. Mustard. 2007. cross-scalar satellite phenology from ground, landsat, and ModiS data. Remote Sensing of Environment 109: 261?273. gu, l., P. J. hanson, W. Mac Post, d. P. kaiser, B. yang, R. nemani, S. g. Pallardy et al. 2008. The 2007 eastern uS spring freezes: increased cold damage in a warming world? Bioscience 58: 253?262. hunter, a. F., and M. J. lechowicz. 1992. Predicting the timing of budburst in temperate trees. Journal of Applied Ecology 29: 597?604. ibanez, i., R. B. Primack, a. J. Miller-Rushing, e. ellwood, h. higuchi, S. d. lee, h. kobori et al. 2010. Forecasting phenology under global warming. Philosophical Transactions of the Royal Society B-Biological Sciences 365: 3247?3260. korner, c., and d. Basler. 2010. Phenology under global warming. Science 327: 1461?1462. lechowicz, M. J. 1984. Why do temperate deciduous trees leaf out at different times? adaptations and ecology of forest communities. American Naturalist 124: 821?842. Menzel, a. 2000. Trends in phenological phases in europe between 1951 and 1996. International Journal of Biometeorology 44: 76?81. Miller-Rushing, a. J., T. l. lloyd-evans, R. B. Primack, and P. Satzinger. 2008. Bird migration times, climate change, and changing population sizes. Global Change Biology 14: 1959?1972. Miller-Rushing, a. J., and R. B. Primack. 2008. global warming and flowering times in Thoreau's concord: a community perspective. Ecology 89: 332?341. Morin, X., M. J. lechowicz, c. augspurger, J. o' keefe, d. Viner, and i. chuine. 2009. leaf phenology in 22 north american tree species during the 21st century. Global Change Biology 15: 961?975. o'keefe J. 2010. Phenology of Woody Species. harvard Forest data archive: hF003. Perry, T. o. 1971. dormancy of trees in winter. Science 171: 29?36. Richardson, a. d., a. S. Bailey, e. g. denny, c. W. Martin, and J. o'keefe. 2006. Phenology of a northern hardwood forest canopy. Global Change Biology 12: 1174?1188. Richardson, a. d., B. h. Braswell, d. y. hollinger, J. P. Jenkins, and S. V. ollinger. 2009a. near-surface remote sensing of spatial and temporal variation in canopy phenology. Ecological Applications 19: 1417?1428. Richardson, a. d., d. y. hollinger, d. B. dail, J. T. lee, J. W. Munger, and J. o'keefe. 2009b. influence of spring phenology on seasonal and annual carbon balance in two contrasting new england forests. Tree Physiology 29: 321?331. Willis, c. g., B. R. Ruhfel, R. B. Primack, a. J. MillerRushing, J. B. losos, and c. c. davis. 2010. Favorable climate change Response explains non-native Species' Success in Thoreau's Woods. PLoS ONE 5(1): e8878. doi:10.1371\/ journal.pone.0008878. caroline Polgar is a graduate student at Boston university, where Richard Primack is a professor. For the past eight years, Richard Primack and his students have been investigating the impact of climate change on the plants and animals of Massachusetts, with much of the focus at the arnold arboretum and concord. "},{"has_event_date":0,"type":"arnoldia","title":"Not-So-Traditional Chinese Medicine: The Example of Donglingcao (Isodon rubescens)","article_sequence":3,"start_page":23,"end_page":30,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25511","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed0708926.jpg","volume":68,"issue_number":4,"year":2011,"series":null,"season":null,"authors":"Harris, Eric S. J.","article_content":"Not-So-Traditional Chinese Medicine: The Example of Donglingcao (Isodon rubescens) Eric S. J. Harris T he use of plants in Traditional Chinese Medicine (TCM) has a long and well-recorded history. Not all plants in the TCM pharmacopoeia share this long history of use, however, and TCM, like any institution of knowledge, continues to grow, expand, and change. As an example, the medicines sold in TCM stores in Chinatowns in the United States include a mix of herbs that have been known for centuries, and some that have been introduced to the TCM pharmacopoeia only in the last few decades. This article provides the context for my research on one of those recent introductions to the TCM pharmacopoeia: an unassuming yet potentially medicinally powerful herb in the mint family called donglingcao. Folk Medicines and Formal TCM One of the earliest written A statue of Shennong (\"The Divine Farmer\") on the campus of the Shanghai Univerworks about botanical medi- sity of Traditional Chinese Medicine. (all photos by the author) cines in TCM is the Divine Farmer's Materia Medica (Shen Nong Ben Cao herbs listed in the Divine Farmer's Materia Jing) which was written over 2,000 years ago Medica remain in use. For example, the book and catalogues more than 300 types of mediincludes herbs such as ginseng (Panax ginseng), cines, most of which are plants (Yang 1998). licorice (Glycyrrhiza uralensis), and goji berries The book has been attributed to the mythical (Lycium chinense). Shennong (the Divine Farmer) who is claimed While works such as the Divine Farmer's to have tasted different plants himself to deterMateria Medica maintain an influence in formine which were poisonous and which could mal TCM practice, the use of different botanibe used as medicine. Many, if not all, of the cal medicines in China continues to transform 24 Arnoldia 68\/4 and grow. For example, the recently published multi-volume work of the Chinese Materia Medica (Ministry of State Administration of Traditional Chinese Medicine 1999) lists more than 9,000 different medicines, or roughly 30 times as many substances as listed in the Divine Farmer's Materia Medica. The Chinese Materia Medica contains different substances, such as animals and minerals, but plants make up the overwhelming majority. The Chinese Materia Medica includes plants that have been in recorded use in China for hundreds of years, and also those that were incorporated only very recently. Of the latter, a large number of the newly recorded botanical medicines were catalogued for the first time in the 1960s and 1970s under Chairman Mao's directions to develop and improve medicine in the Chinese countryside. One result of those efforts was the documentation of plants used as folk medicines (in Chinese, caoyao) (Harris and Yang 2009). Folk botanical medicines are typically used in a limited area in China (e.g., within one province), are generally not accompanied by A pharmacy in a Traditional Chinese Medicine hospital in Beijing. Each wooden drawer holds 2 to 3 different medicines, most of which are plants. Traditional Chinese Medicine 25 Native range of Isodon rubescens by province (shaded in dark green), according to the Flora of China (Li and Hedges 1994). Region of traditional medicinal use of I. rubescens circled in yellow dashed line. Collection sites from field trip in 2009 indicated by red triangles. Yellow River shown in blue. written documentation, and information about their medicinal uses is passed down orally from generation to generation. Folk botanical medicines can be distinguished from formal TCM herbs (in Chinese, zhongyao). Formal TCM herbs typically have a much longer written history, standards for their production and use, are often cultivated, and are well known throughout China. Ginseng, licorice, and goji berries are good examples of these. In general, formal TCM herbs are listed in the official Pharmacopoeia of the People's Republic of China (Chinese Pharmacopoeia Commission 2005). By contrast, folk medicines are not standardized and are usually collected from the wild. There are many exam- ples of folk medicines from throughout China, but one of recent note is the herb donglingcao (Isodon rubescens). The Story of Donglingcao Donglingcao (Isodon rubescens) is a mint family (Lamiaceae) plant within the basils and allies group (Tribe Ocimeae) and is closely related to plants like lavender (Lavandula) and coleus (Solenostemon). The genus Isodon includes about 100 species, most of which occur in Asia, with a few species in Africa (Li and Hedges 1994). Isodon rubescens is distributed throughout central China, usually in dry areas on slopes or in thickets along streams. The area of tradi- 26 Arnoldia 68\/4 Clockwise from upper left: Images of Isodon rubescens--a plant growing in typical habitat, a group of plants in flower, close-up view of flowers, inflorescence structure. tional medicinal use of the plant is in the Taihang Mountain range near the Yellow River in Northern Henan province. Plants of I. rubescens are shrubs that can grow about 1 meter (3.3 feet) tall. They typically have many sprawling branches with ovate leaves (Li and Hedges 1994). Isodon rubescens flowers from late summer into autumn with inflorescences of small (about 1 cm[.4 inches] long) white or purplish flowers. Interestingly, populations of I. rubescens growing in parts of China that experience subfreezing temperatures can sometimes produce ribbons of ice--known as \"ice flowers\"--from the stem (Means 2005). In fact, the Chinese name donglingcao ( ) roughly translates Traditional Chinese Medicine 27 as \"winter-ice herb,\" probably in reference to this phenomenon. Based on the tolerance of the plant to low temperatures, it is possible that donglingcao would be able to grow in gardens in New England. I have not seen it grown here yet, but curious gardeners who would like to try growing it may find seeds of I. rubescens available commercially from some distributors in the United States. The first written records of the medicinal use of donglingcao in China are from the 1960s and 1970s during the period of documentation of China's folk medicines. The plant has been traditionally used for indications such as sore throat and stomach problems. Donglingcao is usually collected in the wild, although there have been some recent efforts at cultivation in northern Henan province. To prepare donglingcao as a medicine, the aerial portions (i.e., stems and leaves) are collected in July and August and dried in the sun. When needed, the dried plant is usually steeped in water to make a tea. The taste of the tea is extremely bitter, owing to diterpenoid chemicals produced by the plant (more about those below). In fact, it is very easy to identify plants of Isodon in the field by simply tasting a leaf-- if after several bites the bitter taste compels you to spit it out, then the plant is likely a species of Isodon. In addition to being taken as a tea, donglingcao has been combined with other herbs in some Chinese patent or proprietary medicines that are available in pill form. During the effort to study folk botanical medicines in the 1970s, it was discovered that one of the bitter diterpenoids in donglingcao, a chemical called oridonin, might have some use in treating cancer (Sun et al. 2006); interestingly, parallel work in Japan on related species of Isodon, such as I. japonicus also came to similar results. This discovery followed the general integration of scientific and chemical approaches in the research of Chinese herbs (in what might be called a \"re-tooling\" of Shennong's approach). A Donglingcao cultivation site in northern Henan province. The plants shown are usually larger in the wild, but this picture was taken in late October after the plants had already been harvested. 28 Arnoldia 68\/4 Commercial products that contain donglingcao include (left to right) throat lozenges, pills, and tea. initial studies in the 1970s. However, in the last few years oridonin has again garnered scientific interest for its potential as an anti-cancer remedy (Zhou et al. 2007). Oridonin, one of the main chemical ingredients responsible for the bioactivity of donglingcao. famous example of the fruits of these efforts is the anti-malarial compound artemisinin. Artemisinin was originally discovered in the 1970s by Chinese researchers examining Artemisia annua, a plant long known in TCM (Hsu 2006). Artemisinin is now used globally as a standard treatment for malaria, and cultivated plants of Artemisia annua remain the main source of the compound today. In contrast to the more popular Artemisia, donglingcao and the chemical oridonin gradually lost the spotlight after the Current Research Starting in 2009, I became interested in donglingcao as an example of a folk botanical medicine that had clear potential to become a standardized TCM herb with more common and widespread use. I began research on this plant to understand the evolution and ecology of the chemicals that are responsible for the medicinal effect, in particular the compound oridonin. The primary goal of this research was to suggest populations or related species of this plant that would have the highest production of oridonin. In collaboration with researchers in the colleges of Chinese medicine in Beijing and Henan province, I traveled to China in the fall of 2009 to collect plants of I. rubescens and related species. The trip included collection areas in Henan province where donglingcao is traditionally used and also other areas in China where I. rubescens is known to occur (Hubei and Guizhou provinces). Through the course of the collecting trip, I traveled together with a Chinese graduate stu- Traditional Chinese Medicine 29 dent by car, sleeper-train, bus, and taxi, covering a distance of more than 1,500 kilometers (over 900 miles) from the northernmost collection site to the southernmost site (roughly the same distance as between Boston and Chicago). This distance allowed me to visit various scenic rural areas, from the expansive Taihang Mountains to small ethnic minority villages in Guizhou province. The collection trip also provided the opportunity to sample a wide variety of Chinese local cooking, from mutton noodle soup in Environment near Isodon collection site in the Taihang mountains, Henan province. Henan in the north to fried \"field chicken\" (=frog) in the south. At each collection site, I collected several individual plants of Isodon to account for possible variability of chemical production within populations. For each individual plant, I collected a pressed voucher specimen, dried leaves in silica desiccant, and a seed sample if the plant was in fruit. The vouchers will be accessioned and deposited at the Harvard University Herbaria. The dried tissue in desiccant has been used for DNA and chemical analyses. The Environment near Isodon collection site in Jiangkou county, Guizhou province. seeds were collected so that chemical content of all populations could be of my work will provide insight into the degree compared in greenhouse grown samples in order of variability in oridonin production in I. rubeto factor out possible differences in chemical scens and related species, and will ideally help production that result from different growing in the standardization of the use of the plant for environments. I have completed the chemical more widespread use in TCM and elsewhere. In and genetic lab work on the wild-collected and fact, although previous editions of the Pharmagreenhouse-grown samples and am now eagerly copoeia of the People's Republic of China did examining and interpreting the results. not list donglingcao, the most recent edition In addition to my primary goal of locating released in the summer of 2010 now includes sources of high oridonin production, the results this plant. It is likely, then, that research and 30 Arnoldia 68\/4 The author in a thicket of Isodon rubescens in Hubei province. development of this plant will continue. With roots as a folk medicine in China, donglingcao has seemingly finally earned its place in the canons of the official Traditional Chinese Medicine. And, depending on the plant's hardiness and desirability as an ornamental, it may also earn its place in gardens in New England. [NOTE: Some of the greenhouse-grown Isodon rubescens plants from this collection trip will be planted at the Arnold Arboretum within the next year or two.] References Chinese Pharmacopoeia Commission. 2005. Pharmacopoeia of the People's Republic of China: 2005. People's Medical Publishing House, Beijing. Harris, E. and B. Yang. 2009. Variation and Standardization in the Use of a Chinese Medicinal Moss. Economic Botany 63: 190?203. Hsu, E. 2006. Reflections on the \"discovery\" of the antimalarial qinghao. British Journal Of Clinical Pharmacology 61: 666?670. Li, X.W. and I.C. Hedges. 1994. Isodon. In: Flora of China, Vol. 17, 269?291. Science Press & Missouri Botanical Garden, Beijing & St. Louis. Means, B. 2005. Mysterious Ice \"Flowers\". The American Gardener Jan\/Feb: 34?37. Ministry of State Administration of Traditional Chinese Medicine. 1999. Chinese Materia Medica. Shanghai Science and Technology Press, Shanghai. In Chinese. Sun, H., S. Huang, and Q. Han. 2006. Diterpenoids from Isodon species and their biological activities. Natural Product Reports 23: 673?698. Yang, S. 1998. The divine farmer's materia medica : a translation of the Shen Nong Ben Cao Jing. 1st ed. Blue Poppy Press, Boulder CO. Zhou, G., S. Chen, Z. Wang, and Z. Chen. 2007. Back to the future of oridonin: again, compound from medicinal herb shows potent antileukemia efficacies in vitro and in vivo. Cell Research 17: 274?276. Eric S. J. Harris, PhD, is a Research Associate at the Harvard University Herbaria. "},{"has_event_date":0,"type":"arnoldia","title":"2010 Weather at the Arboretum","article_sequence":4,"start_page":31,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25509","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed0708528.jpg","volume":68,"issue_number":4,"year":2011,"series":null,"season":null,"authors":"Famiglietti, Bob","article_content":"2010 Weather at the Arboretum Bob Famiglietti A s in 2008 and 2009, above average rainfall continued in 2010. Moisture combined with warm temperatures and plentiful sunshine created optimum growing conditions for the Arboretum's plant collection. January produced less than 6 inches of snow and was relatively mild. The average temperature was almost 5?F warmer than the previous January. A January thaw occurred on the 24th through the 28th, reaching the monthly high temperature of 57?F on the 26th. It dropped to single digits on only three nights, a rare occurrence, reaching a low of 5?F each time. The first Arctic cold front of the year passed through on the 29th with wind gusts up to around 50 mph. The 6-inch snowpack dwindled to a trace by month's end. February was mild and had 4.74 inches of precipitation, much of it falling as rain. The 24th, 25th, and 26th produced strong east winds and 3.88 inches of rain. A low of 13?F was recorded on the 7th, one of the highest lows ever recorded for February. The remainder of the month was unseasonably warm and a high of 49?F was reached on the 21st. Snowfall was light with only 5 inches recorded from two storms. A minimal snowpack created optimum ground conditions, enabling our horticulture staff to perform vital winter pruning of the Arboretum's plants. There was only a trace of snow on the ground by the end of the month. March continued the 2010 pattern of mild temperatures. There was only 0.4 inch of snow for the month. Spring appeared to be on a very early track and, as good gardeners, we hoped for the arrival of abundant spring rains. By month's end--and nearly 18 inches of rain later--we regretted what we had wished for. March started mild with strong winds, rain, and a trace of snow. Temperatures raced into the 50s for five days by mid-month. A three-day Nor'easter produced 3.94 inches of rain on the 14th, 3.74 inches on the 15th, and 1.27 inches on the 16th. A total of 9.43 inches of rain had fallen by midmonth. The rain continued: another 1.51 inches on the 23rd, 0.85 inches on the 24th, with smaller amounts from the 26th through the 29th. The third large storm of the month arrived on the 30th, dropping 3.15 inches that day and 2.15 inches on the 31st, ending the month with an amazing 17.44 inches of precipitation. This was about 13 inches above normal. There were 16 days with measurable precipitation in our rain gauge. March became the wettest month measured at the Arboretum since the weather station was established at the Dana Greenhouses in 1962. (By comparison, the wettest month ever recorded for Boston's official weather site was 17.09 inches in August 1955.) All the rain caused numerous problems throughout the Arboretum: ponds overflowed their banks and flooded adjoining roads and collections, the linden March 15th--heavy rainfall caused bussey brook to overflow, leading to serious erosion in several locations including this spot at the base of hemlock hill near the South Street Gate. (matt connelly) (Tilia) collection looked like a large lake, and Bussey and Goldsmith Brooks overflowed their banks, causing severe erosion. Our secondary road system was severely eroded--90% of it had to be regraded and resurfaced, and additional water mitigation channels had to be installed. In early April a sinkhole was discovered in the road surface over Goldsmith Brook at the Arboretum's main entrance, likely caused when March floodwaters deteriorated the underlying culvert. The entrance was closed to vehicle traffic for nearly a year but re-opened on April 1, 2011. april started warm, reaching 78?F on the 3rd. It reached 90?F on the 7th, breaking a record as the earliest 90?F day in April. A low of 32?F occurred on the 28th. It was the only freezing temperature recorded for the month-- a rare event--and the last frost date for the season. April ended on the dry side with no snow and only 2.13 inches of rain. This gave our nurseries a chance to dry out so our spring transplanting season could begin. May was warm, dry, and sunny. A high of 94?F on the 26th and one of the sunniest Mays on record helped push the average high temperature over 4?F above normal, ranking it the 5th warmest on record. The first five months of 2010 averaged over 3?F above normal. Precipitation was below normal at 2.92 inches. Lilac Sunday on May 9th was partly sunny but cool and very windy. Visitors walked along a narrow dry strip on Forest hills road on March 16th, with an overflowing Dawson pond in the background. (kevin b. schofield) arboretum horticultural technologist Kit Ganshaw measures water depth in rehder pond on March 25th after yet more rain fell on the previous two days. (nancy rose) Weather 33 Arnold Arboretum Weather Station Data ? 2010 avg. Max. (?F) Jan Feb Mar apr May Jun Jul auG Sep OcT nOV Dec 35.2 37.2 50.3 61.6 73.1 79.6 86.2 81.4 77.2 62.8 51.5 37.5 avg. Min. (?F) 20.1 24.3 35.2 42.1 52.0 60.7 68.0 62.7 57.1 44.0 34.0 23.4 avg. Temp. (?F) 27.7 30.8 42.8 51.9 62.6 70.2 77.1 72.1 67.2 53.4 42.8 30.5 Max. Temp. (?F) 57 49 72 90 94 93 98 92 95 82 65 56 Min. Temp. (?F) 5 5 24 32 38 49 54 52 45 33 25 12 precipi- Snowtation fall (inches) (inches) 3.74 4.74 17.44 2.13 2.92 3.52 2.63 8.05 1.99 5.42 3.75 3.93 16.1 5.3 5.0 .4 average Maximum Temperature . . . . . . . . . . 61.1?F average Minimum Temperature . . . . . . . . . . 43.6?F average Temperature . . . . . . . . . . . . . . . . . . . 52.4?F Total precipitation . . . . . . . . . . . . . . . . . . . . . 60.26 inches Total Snowfall. . . . . . . . . . . . . . . . . . . . . . . . . 26.8 inches Warmest Temperature . . . . . . . . . . . . . . . . . . 98?F on July 7 coldest Temperature . . . . . . . . . . . . . . . . . . . 5?F on January 5 and February 1 last Frost Date . . . . . . . . . . . . . . . . . . . . . . . . 32?F on april 28 First Frost Date . . . . . . . . . . . . . . . . . . . . . . . . 29?F on november 1 Growing Season . . . . . . . . . . . . . . . . . . . . . . . 186 days 34 Arnoldia 68\/4 June continued very warm, reaching 93?F on the 24th and 28th. It was the 15th warmest June in Boston's 139 years of weather record keeping. In contrast, June 2009 was the 3rd coldest. There were 17 days of measurable rain but only 3.52 inches fell for the month. Thunder was detected on six days, double the average. July was very hot and somewhat dry. The average high temperature was 86.2?F, making it the 3rd hottest July on record. It reached 100?F at Logan Airport in Boston for the first time since 2002. It hit at least 90?F twelve times this month alone, marking 17 days already for this year, more than the entire annual average for Boston. We had three heat waves (highs of at least 90?F for 3 consecutive days) this month. An Arboretum high of 98?F was reached on the 6th. In contrast, last June?July was the 4th coldest in the 138 years of Boston's weather records. Rain totaled 2.63 inches, in contrast to last July's nearly 8 inches. auGuST kept pace with the preceding months as warmer than normal, though not as extreme as July. A high of 92?F occurred on the 9th. This summer (June?August) was the third warmest on record. It was also a wet month as 8.05 inches of rain fell, though rainfall occurred on only eight days. 6.43 inches of rain fell on the 22nd through the 26th. Three 90?F days ended the month as another heat wave began. SepTeMber also was very warm and dry with only 1.99 inches of rain. Our only thunderstorm occurred on the 8th. The month started as August had ended with temperatures in the 90s for the first three days. A high for the month of 95?F was reached on the 2nd. This was the third warmest September on record. Wind gusted to nearly 50 mph on the 30th as Tropical Storm Nicole passed offshore. OcTOber was on track with the preceding 9 months as we continued with above average temperatures. A high of 82?F was reached on the 1st, our last reading in the 80s for the year. Decent rainfall occurred with a total of 5.42 inches recorded. Precipitation was recorded on 14 days but the remaining ones were often sunny and glorious. There was no trace of snow this month but a low of 33?F was recorded on the 22nd, bringing just a hint of light frost in the low areas around the Arboretum ponds. This spectacular month of weather made for grand viewing of our fall foliage. nOVeMber became our first colder than normal month of 2010. The Arboretum's first freeze occurred the night of the 1st when temperatures dropped to 29?F, ending our growing season. A heavy widespread frost occurred on the 3rd. Rainfall was near normal with 3.75 inches. There was no trace of snow this month, and a high of 65?F was recorded on the 17th. Finally as temperatures cooled we sensed the impending dormant season, a prerequisite to winter storage of our containerized plant material from the Dana Greenhouses. Weather 35 Glowing orange and gold sugar maple (Acer saccharum) foliage framed the leventritt Shrub and Vine Garden on October 25th. (michael dosmann) DeceMber was snowy and the only month of 2010 with markedly below normal temperatures. A low of 12?F was reached on the 10th and then a high of 56?F was recorded on the 13th, an extreme range of 44?F in three days. Our first snowfall occurred on the 21st, leaving 2 inches on the ground. There was a large, violent Nor'easter on the 26th and 27th. Over a foot of snow was deposited but blizzard-level wind conditions created snowdrifts 3 to 4 feet deep, making it difficult to measure exactly how much snow fell. This fierce storm left some of the Arboretum's plants with structural damage. The year ended with a snow depth of nearly 1 foot, a sign of things to come. Bob Famiglietti is a Horticultural Technologist at the Arnold Arboretum's Dana Greenhouses. "},{"has_event_date":0,"type":"arnoldia","title":"The Family Tree: Prunus 'Hally Jolivette'","article_sequence":5,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25513","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4e1d260a36f.jpg","volume":68,"issue_number":4,"year":2011,"series":null,"season":null,"authors":"Sax, Miles S.","article_content":"The Family Tree: Prunus `Hally Jolivette' Miles Sax A sk someone to show you their family tree and you'll likely be shown a genealogy chart in an old book, or perhaps a family lineage document on the computer. As a child, if I asked my father to see our family tree he would take my hand, lead me outside, and show me the enchanting spring blooms of the `Hally Jolivette' cherry in our yard. This tree-- named for my great-grandmother--is important to me both as a link to my family history and as a horticultural gem. Prunus `Hally Jolivette' is a flowering cherry hybridized by Dr. Karl Sax, the Arnold Arboretum's fifth director (and my great-grandfather). A research scientist by training, Sax's investigations in genetics and chromosome studies played a seminal role in the biological sciences of his time, most notably in the field of cytology. His interests also encompassed plant breeding, a field to which he made many contributions through his extensive hybridizing studies. Conducting much of his work while a professor at the neighboring Bussey Institute, many of Sax's hybrids \"jumped the fence\" and ended up in the Arboretum's collections, and a number were introduced to the nursery trade. Perhaps the best of Sax's hybrids, Prunus `Hally Jolivette' was introduced by the Arnold Arboretum in 1948. It resulted from crossing P. subhirtella and P. x yedoensis, and then backcrossing with P. subhirtella. It is a fine-textured, densely branched, rounded small tree or large shrub that grows about 15 feet (4.6 meters) tall and equally wide. Each spring it delights the eye with a profusion of pink buds opening up to 1? inch (3.2 centimeter) diameter pinkish white double flowers. At the Arboretum, flowering occurs in late April or early May. Bloom may continue for ten to twenty days, a notably long period for a flowering cherry. Flowers open prior to leaf emergence so the floral effect is unobscured. The 2 to 3 inch (5.1 to 7.6 centimeter) long leaves are simple, alternate, and dark green. `Hally Jolivette' cherry grows best in full sun, is drought tolerant, and is an ideal plant for small gardens, specimen plantings, or even bonsai. Best adapted to USDA zones 5 through 7, this cherry is a precocious bloomer and often flowers in its second year of growth. Three young specimens (accessions 278-2007-B, C, and D) can be seen in the Arboretum's newly renovated Bradley Rosaceous Collection. Prunus `Hally Jolivette' has received many accolades including awards from both the Pennsylvania Horticultural Society and Cornell Cooperative Extension. Of his many introductions, Karl Sax clearly held this Prunus in a special place because he named it in honor of his wife, colleague, and scientist in her own right, Dr. Hally Jolivette. She was a mycologist and botanist, and in 1912 was the first woman to receive a Ph.D from Stanford University in the field of botany. She held many academic posts across the country including instructorships at Washington State University, Wellesley College, and the Bussey Institute. She was a research scientist and published original work as well as co-authoring many papers with Dr. Karl Sax. Both a scientist and a devoted mother, she is remembered as a woman who put cookies on the table for her three children one minute and was deep in study with her microscope the next. My family has been honored and delighted to have this tree that not only greets each spring with great beauty but also reminds us of our lineage. In the words of plantsman Michael Dirr \"Great plants transcend the generations,\" and with this in mind I hope that Prunus `Hally Jolivette' will be planted and enjoyed for many years to come. Miles Sax is a horticultural apprentice at the Arnold Arboretum, where he works on evaluating, maintaining, and improving the Malus collection. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23424","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160bb28.jpg","title":"2011-68-4","volume":68,"issue_number":4,"year":2011,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"White Bracts of the Dove Tree (Davidia involucrata): Umbrella and Pollinator Lure?","article_sequence":1,"start_page":2,"end_page":10,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25508","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070816f.jpg","volume":68,"issue_number":3,"year":2011,"series":null,"season":null,"authors":"Sun, Ji-Fan; Huang, Shuang-Quan","article_content":"White Bracts of the Dove Tree (Davidia involucrata): Umbrella and Pollinator Lure? Ji-Fan Sun and Shuang-Quan Huang JI-FAN SuN AND SHuANg-QuAN HuANg P ollen movement in flowering plants depends on various vectors including animals, wind, and water. Compared to wind- or waterpollinated flowers, animal-pollinated flowers are generally showier, often with bright colors. They also often produce nectar or other rewards to attract pollinators. However, the advertisement and reward for pollinators may also attract plant enemies. Herbivores can consume parts of flower structures, entire flowers, or whole plants. For example, nectar robbers may penetrate a hole in the corolla and thereby suck nectar from flowers without playing a pollination role. The great diversity found in angiosperm flowers can be fully understood only when the diverse floral traits are considered as functional units, shaped by partly opposing selective pressures (Faegri and van der Pijl 1979; Waser and Ollerton 2006). In experimental work on the evolution of floral traits, most attention has centered on natural selection that favors mutualistic pollinators and hinders antagonistic herbivores (Fenster et al. 2004; Strauss and Whittall 2006). A Closer Look at Dove Tree The dove tree, Davidia involucrata, A dove tree in bloom at the study site in Shennongjia Nature Reserve, is a species prized by gardeners for its Hubei, China. showy bracts. The only species in the genus, dove tree is a medium-sized tree (up was more widespread in the past (for a review, to 20 meters [65 feet] tall) that is endemic in see Manchester 2003). mountain forests at altitudes of 1,100 to 2,600 The genus Davidia is named after Father meters (3,600 to 8,500 feet) in western China Armand David (18261900), a French mission(Fang and Chang 1983). Fossils from the Paleoary and keen naturalist who lived in China from cene of North America indicate that the lineage 1862 to 1874 and collected many specimens of White Bracts of the Dove Tree 3 PAuL MEyER PETER DEL TREDICI Dove trees now grow in gardens around the world, including these specimens at the Morris Arboretum in Philadelphia, Pennsylvania (left), and the VanDusen Botanical Garden in Vancouver, British Columbia (right). plants previously unknown in the West (Zhang and Li 1994). David was also the first westerner to describe another rare Chinese endemic, the giant panda (Ailuropoda melanoleuca). The dove tree has been considered a firstclass endangered plant in China, but it became well-known in its homeland only after it was seen growing in other countries during two diplomatic visits by Chinese leaders. When Enlai Zhou (18981976), the first premier of the People's Republic of China, visited geneva, Switzerland, in 1954, he was impressed by the beauty of flowering dove trees in the gardens of many local families and was told that the tree came from China. Another surprise happened when Chinese leaders visiting Washington, D.C., saw dove trees in bloom in front of the White House in the early 1970s. The tree's common names--dove tree or handkerchief tree--refer to the two white, paperlike bracts that surround the base of each flower head (capitulum). The bracts initially are small and green, resembling leaves, but increase in size and turn white as the flowers mature. The change in bract color from green to white is associated with the bracts becoming uV (ultraviolet)-light-absorbing (Burr and Barthlott 1993). The anthers are even more strongly uVlight-absorbing. This trait is associated with attracting pollinating insects that see uV light, and the species has therefore been classified as entomophilous (insect pollinated) (Burr and Barthlott 1993), although the pollination of Davidia involucrata had not been previously studied in the field. We are particularly interested in the questions \"What is the function of the white bracts? Do the dove tree's bracts play a role in attracting pollinators?\" Many plants within Cornales, the dogwood clade, have large bracts that surround the inflo- 4 Arnoldia 68\/3 COuRTESy OF KEVIN NIxON flowers in a population of 16 flowering dove trees to assess the function of this species' bracts. Though dove tree's mature bracts draw much attention, the round inflorescences dangling beneath the bracts are also interesting. The dark purple inflorescences generally consist of a single perfect (holding both stamens and pistils) flower surrounded by numerous male flowers (Fang and Chang 1983). The individual flowers are naked and nectarless, without The colorful bracts surrounding the small flowers of this Dalechampia attract pollinators. sepals or petals. Before rescences. Bracts have been thought to protect flowering, dove tree's bracts are green, turning flowers from herbivores in various species. white rapidly as flowers mature and anthers For example, the bracts of Dalechampia vines begin to dehisce. The anthers split longitufunction as honest signals to pollinators of the dinally, and pollen grains are exposed on the presence of floral reward (Armbruster et al. recurved anther walls. Individual capitula of the 2005) and also function in protection from dove tree last 5 to 7 days and the bracts drop off florivores (flower eaters) and pollen thieves when flowering ends. (Armbruster 1997). As part of the study, we collected both green Flowers are also under strong selection presand white bracts and preserved them for later sures from their physical environment, yet observation under the microscope. We also research on selection by abiotic environmenmeasured the length and width of the bracts on tal factors on flowers has been limited (Corbet each of 20 capitula daily between April 16 and 1990; galen 2005). During our previous field April 22, 2006. work, our observation was often interrupted by Do Dove Tree's Bracts Function in the rainy days. Rain is one of those abiotic factors Pollination Process? that acts as a selective agent on flowers. This To investigate the role of bracts in pollinator was first recognized by Sprengel ([1793]1972), attraction, we recorded pollinator visits to four who noted that rain may wash away pollen kinds of capitula: (1) natural, (2) with both bracts grains and dilute flower nectar. This added removed, (3) with both bracts replaced by green another question to our research: Did the need artificial bracts made of copy paper, or (4) with to protect pollen grains from rain play a role in both bracts replaced by white artificial bracts the evolution of large bracts? made of copy paper. The artificial bracts were Carrying Out the Study similar in shape and size to the natural bracts. With our questions about rain protection and We recorded the number of pollinator visits to pollinator attraction in mind, we carried out the dove trees' flowers from 10:00 in the mornobservations and experiments in a natural poping to 3:00 in the afternoon between April 18 ulation of dove trees in western China (Shenand April 22 in 2005, and between April 16 nongjia Nature Reserve, Hubei) in April 2005 and April 20 in 2006. Two observers monitored and April 2006. In our study, we manipulated two sites during these periods of maximum pol- White Bracts of the Dove Tree 5 JI-FAN SuN AND SHuANg-QuAN HuANg Clockwise from upper left: A young inflorescence displays still-green bracts and deep purple immature anthers. An artificial inflorescence in which the natural white bracts were replaced with green paper. Two artificial inflorescences in which the natural white bracts were replaced with white paper. Two mature inflorescences, the upper one is untouched, while the lower one has had the bracts removed. linator activity. Insect visits became extremely rare after 3:00 in the afternoon, and during one night of observations, no pollinators were observed visiting this nectarless species. Pollinators were collected and sent to the Institute of Zoology of the Chinese Academy of Science in Beijing for identification. To examine whether capitula with intact bracts lost more or less pollen to rain than did capitula that had their bracts removed, late in the flowering period we counted the pollen remaining within 29 and 27 capitula that were natural or had both bracts removed, respectively, and had experienced at least one rainy day. 6 Arnoldia 68\/3 Numbers of anthers and ovules per capitulum and pollen grains produced per anther were estimated in 34 capitula with freshly dehiscing anthers. In September 2005 and 2006, we collected 39 and 32 open-pollinated capitula and counted their seeds to estimate seed set under natural pollination. What We Found Out... When we looked at the dove tree bracts that we had collected at either the green or white stage, we saw that the parenchyma cells in the green bracts were full of chloroplasts, while the parenchyma cells of white bracts had degenerated and contained few chloroplasts. Bract color turned from green to white on the third or fourth day, when the anthers began to dehisce and when bracts had reached three-quarters of their final size. Bracts continued to grow during anthesis but dropped off soon thereafter. Pollen-collecting bees and pollen-feeding beetles were the major visitors and pollinators of the flowers. We recorded a total of 2,174 visits to capitula, of which bees and beetles accounted for 93.6% and 6.4%, respectively. Beetles generally stayed within a capitulum for 1 to 2 hours, while bees spent 4 to 6 seconds per capitulum foraging for pollen, suggesting that bees are the more effective pollinators of the dove tree. Bees that visited included Apis cerana, Xylocopa appendiculata (Apidae), and Halictus and Lasioglossum species (Halictidae), and beetles included Agriotes species (Elateridae), Oxycetonia jucunda (Cetoniidae), and species of Nitidulidae. Visitor frequencies were low, and so was seed set of openpollinated capitula in both years. It seems clear that wind pollination Pollen grains of Davidia involucrata germinating in 10% sucrose solution. of dove tree is unlikely since its Pollen Stickiness and Behavior in Water The pollen grains of anemophilous (wind pollinated) plants are not sticky, while those of entomophilous plants usually are sticky in order to adhere to visiting pollinators. To assess the possibility of wind pollination of dove tree, we tested the stickiness of its pollen grains by their adherence to glass slides. We also placed netting around 20 capitula, thereby excluding pollinators but allowing possible wind pollination, and later examined the netted capitula for seed production. To test the behavior of pollen grains under rainy conditions, we followed Huang et al.'s (2002) method of pollen germination. Pollen grains from newly dehisced anthers were placed in sucrose solutions of 5%, 10%, 15%, and 20% by mass to examine optimum conditions for pollen germination. We then compared germination rates of pollen grains from eight ran- domly collected capitula in distilled water or in the optimum sucrose solution (10%). Pollen grains that had either germinated or burst after 4 hours were counted under a light microscope. Ten samples of pollen grains from each flower were analyzed in this manner. Data analysis was accomplished by using one-way ANOVA analysis to compare the frequencies of pollinator visits to the four kinds of capitula and the amount of pollen remaining in capitula after different experimental treatments and to assess pollen viability in distilled water versus 10% sucrose solution. JI-FAN SuN AND SHuANg-QuAN HuANg White Bracts of the Dove Tree 7 JI-FAN SuN AND SHuANg-QuAN HuANg Cross sections of a green bract (left) and a white bract (right). Mesophyll cells and chloroplasts have degenerated in the white bract. Scale bars = 0.5 millimeters (0.02 inch). pollen grains are sticky and apparently not picked up by wind. Also, the netted capitula did not produce any seeds, further evidence that insects rather than wind pollinate dove tree flowers. Bees preferred to visit capitula with white bracts over those with green bracts, on the basis of four consecutive days of observations on inflorescences of 62 natural capitula, 62 bractless capitula, 62 white-papered capitula, and 48 green-papered capitula. Visits to natural and white-papered inflorescences were not significantly different but were higher than those in the other two treatments. Visits to inflorescences with bracts removed and those with bracts replaced by green paper did not differ significantly. The pollen\/ovule ratio in D. involucrata is extremely high, with a capitulum producing only about 7 ovules to about 900 to 1,000 anthers, which produced over a million pollen The Asian honey bee (Apis cerana) was one of the pollinators found on dove tree flowers. The specimen seen here is feeding from Chinese elderberry (Sambucus chinensis) growing in Sichuan, China. grains in total. Pollen amounts remaining in capitula with their bracts removed were not significantly different from those in intact capitula but were significantly lower than the ZACHARy HuANg, HTTP:\/\/BEES.MSu.EDu 8 Arnoldia 68\/3 total pollen production, demonstrating high pollen loss to rain or removal by pollinators. Pollen germinated best in 10% sucrose solution, and there was a highly significant difference in germination and bursting rates between pollen grains placed in distilled water or 10% sucrose solution. After 4 hours, about 85% of pollen grains in water had burst, while only about 37% of the grains in 10% sucrose solution had burst. (ed. note: see Sun et al. 2008 for detailed results) ... and What It Means Pollen-collecting bees, the most important pollinators of Davidia involucrata, preferred visiting white-bracted capitula, and it is therefore likely that during their green stage the bracts function in photosynthesis, while during their JI-FAN SuN AND SHuANg-QuAN HuANg white, uV-light-absorbing stage their function changes to attracting pollinators. During more than 170 hours of observation, we never observed bees visiting green-bracted capitula. Their uV-light absorbance makes the white bracts stand out from surrounding foliage in the bee visual spectrum (Burr and Barthlott 1993; Kevan et al. 1996) and is likely due to the flavonoids that are the major pigments in the bracts of D. involucrata (Hu et al. 2007). Over the 2 years of our study, insect visitation was low. With low visitation rates, prolonging the flowering period will benefit reproductive success as a sit-and-wait strategy (Ashman and Schoen 1994). Protection of the pollen grains presented on the recurved anther walls during the 5 to 7 day flowering period would then be of key importance; the longer viable pollen is Greenish immature bracts function photosynthetically, while mature white bracts absorb UV light and function in attracting pollinators. Note that many of the anthers have already dehisced in the mature inflorescences. White Bracts of the Dove Tree 9 ROBERT MAyER present, the better the chance of a pollinator visiting within that period. Since dove tree's flowers lack a corolla, protection of the pollen must be achieved by the bracts. Dove trees generally flower from mid-April to mid-May, a period which is within the rainy season of the subtropical region where they occur. Since the pollen grains of dove tree readily burst in water, it is probable that the rooflike bracts function as an umbrella to reduce rain damage to the anthers and pollen. Indeed, we repeatedly observed rain-damaged stamens in the capitula where we had removed bracts, while capitula with intact bracts had dry stamens even after heavy rain. The fact that bract-bearing capitula lost as much pollen as bractless ones is attributable to bees removing most pollen from the former, while rain washed away most pollen from the latter. The evolution of flower or inflorescence structures with multiple functions may reflect the net effect of conflicting or additive selective pressures (Anderson 1976; Armbruster 1996, 1997, 2001; galen 1999; Fenster et al. 2004; Armbruster et al. 2005; Strauss and Whittall 2006; Waser and Ollerton 2006). In the case of the dove tree, pollinator selection favors white bracts, while the need to protect pollen from rain favors large bracts. Notably, dove tree's two bracts differ in size, perhaps in response to constraints on weight or resources. Experimental studies of the effects of rain on floral traits are scarce (Bynum and Smith 2001; Huang et al. 2002; galen 2005). Recent studies of pollen longevity in 80 angiosperms found that pollen life-span was decreased by direct contact with water (Mao and Huang 2009). Another evolutionary tactic is seen in the erect flowers of Primula vulgaris; though the flowers regularly fill with water, roughly a quarter of the primula's pollen grains will still germinate even after 5 hours underwater (Eisikowitch and Woodell 1975). While some species have relatively high water-resistant pollen, others rely on nodding flowers, or close their flowers by petal movements (Bynum and Smith 2001; Hase et al. 2006; He et al. 2006), or have other features protecting pollen from wetting, such as in the dove tree. The pale green bracts above these Tilia (linden) flowers may serve several purposes, including attracting pollinators. It seems that the bracts of the dove tree are a striking example of the multiple roles played by one structure in photosynthesis, pollinator attraction, and as a stamen rain shelter. As another example, in Tilia the bracts not only help wind dispersal of the fruits once they mature, but also may act as flags that attract pollinators-- especially nocturnal pollinators-- to the flowers (Anderson 1976). Our observation of the multifunctional nature of bracts in the dove tree suggested that floral traits are under selection pressure from mutualists and antagonists as well as selection by abiotic environmental factors. Acknowledgments This public introduction of our work in the dove tree is based on the first author's PhD thesis and a paper in American Naturalist (2008). We thank y.-B. gong, Q. Fang, and S. S. Renner for collaboration, and we also thank L. Wang from the Altar Park in the Shennongjia Nature Reserve as well as W. Chen, Q. Li, y.-y. Mao, and M.-Q. Pan from Wuhan university for help in the field and in the lab, and S. Armbruster and an anonymous reviewer for many valuable suggestions for an early draft. A grant from the National Science Foundation of China (30825005) to S.-Q. Huang supported this work. Literature Cited Anderson, g. J. 1976. The pollination biology of Tilia. American Journal of Botany 63: 12031212. Armbruster, W. S. 1996. Evolution of floral morphology and function: an integrative approach to 10 Arnoldia 68\/3 adaptation, constraint, and compromise in Dalechampia (Euphorbiaceae). In: D. g. Lloyd and S. C. H. Barrett, eds. Floral Biology, pp. 241272. Chapman & Hall, New york. Armbruster, W. S. 1997. Exaptations link the evolution of plant-herbivore and plant-pollinator interactions: a phylogenetic inquiry. Ecology 78: 16611674. Armbruster, W. S. 2001. Evolution of floral form: electrostatic forces, pollination, and adaptive compromise. New Phytologist 152: 181183. Armbruster, W. S., L. Antonsen, and C. Pelabon. 2005. Phenotypic selection on Dalechampia blossoms: honest signaling affects pollination success. Ecology 86: 33233333. Ashman, T. L., and D. J. Schoen. 1994. How long should flowers live? Nature 371: 788791. Burr, B., and W. Barthlott. 1993. untersuchungen zur ultraviolettreflexion von Angiospermenbluten. II. Magnoliidae, Ranunculidae, Hamamelididae, Caryophyllidae, Rosidae. Trop. subtrop. Pflanzenwelt 87, Akad. Wiss. Lit. Mainz. F. Steiner Verlag, Stuttgart, 193 S. Bynum, M.R., and W. K. Smith. 2001. Floral movements in response to thunderstor ms improve reproductive effort in the alpine species Gentiana algida (gentianaceae). American Journal of Botany 88: 10881095. Corbet, S. A. 1990. Pollination and the weather. Israel Journal of Botany 39: 1330. Eisikowitch, D., and S. R. J. Woodell, 1975. The effect of water on pollen germination in two species of Primula. Evolution 28: 692694. Faegri, K., and L.van der Pijl. 1979. The principles of pollination ecology. 3rd ed. Pergamon Press, Oxford. Fang, W.-P., and C.-y. Chang. 1983. Flora Republicae Popularis Sinicae, vol. 52(2). Pages 157159. Science Press, Beijing. Fenster, C. B., W. S. Armbruster, P. Wilson, R. Dudash, and J. D. Thomson. 2004. Pollination syndromes and floral specialization. Annual Review of Ecology, Evolution and Systematics 35: 375403. galen, C. 1999. Why do flowers vary? The functional ecology of variation in flower size and form within natural plant populations. Bioscience 49: 631640. galen, C. 2005. It never rains but then it pours: the diverse effects of water on flower integrity and function. In: R. Edward and F. A. Bazzaz, eds. Reproductive allocation in plants, pp. 7795. Elsevier Academic Press, Burlington, MA, uSA. Hase, A. V., R. M. Cowling, and A. g. Ellis. 2006. Petal movement in cape wildflowers protects pollen from exposure to moisture. Plant Ecology 184: 587. He, y.-P., y.-W. Duan, J.-Q. Liu, and W. K. Smith. 2006. Floral closure in response to temperature and pollination in Gentiana straminea Maxim. (gentianaceae), an alpine perennial in the Qinghai-Tibetan Plateau. Plant Systematics and Evolution 256: 1733. Hu, J.-y., S.-L. Zhang, Z.-x. Su, and y.-M. Liao. 2007. Pollinator attraction by Davidia involucrata. I. Color. Journal of Plant Ecology 31: 166171. (In Chinese with English Abstract) Huang, S.-Q., y. Takahashi, and A. Dafni. 2002. Why does the flower stalk of Pulsatilla cernua (Ranunculaceae) bend during anthesis? American Journal of Botany 89: 15991603. Jones, C. E. 1967. Some evolutionary aspects of a water stress on flowering in the tropics. Turrialba 17: 188190. Kevan, P., M. giurfa, and L. Chittka. 1996. Why are there so many and so few white flowers. Trends in Plant Science 1: 280284. Mao y.-y., and S.-Q. Huang. 2009. Pollen resistance to water in 80 angiosperm species: flower structures protect rain susceptible pollen. New Phytologist 183: 892899. Manchester, S. R. 2003. Leaves and fruits of Davidia (Cor nales) from the Paleocene of North America. Systematic Botany 27: 368382. Sprengel, C. K. 1793. Das entdeckte Geheimnis der Natur im Bau und in der Befruchtung der Blumen (Reprinted 1972). Weldon & Wesley, New york. Strauss, S. y., and J. B. Whittall. 2006. Non-pollinator agents of selection on floral traits. In: L. D. Harder and S. C. H. Barrett, eds. Ecology and evolution of flowers, pp. 120138. Oxford university Press, Oxford. Sun, J.-F., y.-B. gong, S. S. Renner, and S.-Q. Huang. 2008. Multifunctional bracts in the dove tree Davidia involucrata (Nyssaceae: Cornales): rain protection and pollinator attraction. The American Naturalist 171: 119124 Waser, N. M., and J. Ollerton. 2006. Plant-pollinator interactions: from specialization to generalization. university of Chicago Press, Chicago. Zhang, J., J. Li, and x. Lian. 1994. Morphology and biological characteristics of Davidia involucrata. Journal of Beijing Forestry University 16: 3337. Ji-Fan Sun and Shuang-Quan Huang are from the College of Life Sciences at Wuhan university in Wuhan, China. "},{"has_event_date":0,"type":"arnoldia","title":"Little Big Plant, Box Huckleberry (Gaylussacia brachycera)","article_sequence":2,"start_page":11,"end_page":18,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25507","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070bb6b.jpg","volume":68,"issue_number":3,"year":2011,"series":null,"season":null,"authors":"Nicholson, Robert G.","article_content":"Little Big Plant, Box Huckleberry (Gaylussacia brachycera) Rob Nicholson I t is one of the charms of the science of botany that the most subtle species can contain the most elegant mysteries. Such is the case with box huckleberry (Gaylussacia brachycera), an evergreen subshrub with small, glossy, leathery leaves. Box huckleberry grows in the full shade of mixed pineoak forests in sporadic locations from Pennsylvania to Tennessee (including a fairly recently discovered site in North Carolina [Wilbur 2004]). While this species has been known to botanists for Box huckleberry's foliage looks somewhat similar to boxwood (Buxus spp.), which centuries, it is an example explains its common name. (photo by rob nicholson) of how knowledge builds on knowledge and how even familiar subjects can species found in the hills and mountains near still be a source of discovery. the Argentine border. A few more species are This low-growing shrub is in the Ericaceae Andean, but then a separation of 1,100 miles (heath family), an assemblage of plants that (1,770 kilometers) occurs before reaching any of includes heaths (Erica) and heathers (Calluna), the North American species, the closest being rhododendrons and azaleas (Rhododendron), Gaylussacia dumosa in southern Florida. Eight blueberries and cranberries (Vaccinium), and Gaylussacia species are known from the eastmadrones (Arbutus). It is a fairly large family ern United States and Canada, but none have with species found throughout much of the been found in Mexico, Central America, or the world. The huckleberries, along with the blueCaribbean--a somewhat mysterious gap. berries, are grouped in a subfamily called the The huckleberries have traditionally been Vaccinioideae. Blueberries (Vaccinium) and further divided into three subgroups called sechuckleberries (Gaylussacia) are traditionally tions: section Vitis-idaea, with only Gaylusseparated botanically by the number of chamsacia brachycera; section Decamerium, with bers (called locules) within the fruit and the three North American species; and section size of the seeds. Gaylussacia, with G. mosieri of the Florida While the name huckleberry may bring panhandle and all the South American species. forward associations to Mark Twain's all(Some recent research [Floyd 2002] questions American rascal, Huckleberry Finn, the genus whether Gaylussacia should be divided into Gaylussacia shows a predominantly South sections at all.) So this odd distribution of speAmerican bent. The center of species diversity cies and subgroups presents the first botanifor the genus is southeastern Brazil, with 37 cal puzzle that heavily involves the little box Box Huckleberry 13 Have Berries, Will Travel The migration route traveled by Gaylussacia between the southern and northern landmasses is unclear, with one botanist (Camp 1941) postulating a gradual migration over a former landbridge, a connection to the east of the Isthmus of Panama. Another (Floyd 2002) suggests the possibility that the genus originated in North America, rather than South America. Gaylussacia brachycera is different from all other species within the genus in that it lacks glands upon its leaf surfaces, and is therefore segregated into its own section. Its original discoverer, the French botanist Andre Michaux, thought it to be a Vaccinium and published it as such in 1803. However, it has an ovary split into ten chambers like its brethren Gaylussacia rather than the five chambers usually associated with blueberry, and it also has large seeds rather than the tiny seeds typical of blueberries. Anyone who has sampled wild huckleberries knows they have more crunch than blueberries and are probably less developed as a food crop because of this seediness. I had reason to believe I had found a \"missing link\" on the path between North and South America while collecting in the high pine forests of northeast Mexico. I came upon a lowgrowing, thick-leafed plant that I immediately took to be a species of Gaylussacia because of its close resemblance to box huckleberry. I was excited by the biogeographic implications, and cuttings were collected and brought back to the botanic garden for propagation. When the resulting plants finally flowered I was able to dissect the flowers and determine the plant's true identity. If it was a blueberry then the ovary of the flower would have five compartments, if a huckleberry, then ten. I focused my microscope on the sectioned ovary and saw a pie of five wedges resolve itself. The plant was determined to be a Mexican species of blueberry, Vaccinium kunthianum. It was my ulti- Gaylussacia seeds (seen here) are larger than those of Vaccinium, giving huckleberries a distinct crunchiness when eaten. mate anti-Archimedean moment, the crashing flip side to \"Eureka!\" Depending on which taxonomic interpretation one subscribes to, Gaylussacia brachycera is a unique species among the huckleberries, or has two very close relatives in southern Brazil. Alternative positions have constantly swirled around box huckleberry; Camp (1941) wrote \"it would certainly appear to merit generic rank,\" while recent molecular genetic studies by Dr. Jennifer Whitehead Floyd (2002) show that the box huckleberry may be intermediate between the huckleberries (Gaylussacia) and blueberries (Vaccinium) and may be an ancient hybrid involving species from each camp. But if it is a hybrid, what were the parents? And where might the ancestral lines be? Further molecular genetics studies may finally crack the riddle or even return the species to the Vaccinium fold, where it started with Monsieur Michaux two hundred years ago. How Old Is Old? Gaylussacia brachycera was first found around 1796 by the French botanist Andre Michaux at Warm Springs, Virginia, a locale now shrouded in confusion. Two other collectors found it in the early 1800s, both in West Virginia. Fifty years would pass before another, more northern stand in Perry County, Pennsylvania, was discovered by Spencer F. Baird, a young professor of natural history at Dickinson College who later went on to be Secretary of the Smithsonian Institution. Facing page: Black huckleberry (Gaylussacia baccata) is distributed throughout the eastern United States and eastern Canada. This deciduous Gaylussacia species is noted for its bright red autumn foliage. (photo by nancy rose) RoB NICHolSoN huckleberry: on which continent did the huckleberries first evolve, and how did they then spread? And where did this unique species-- Gaylussacia brachycera--come from? How Old Are You Now? THE MEASUREMENT of age in living plants can be done with a limited number of species. only those ligneous (woody) plants that live in temperate or arctic regions and are exposed to annual weather cycles will dependably create rings that correlate to age. The title holder for oldest plant is still under contention. Among the most vaunted contenders are the Great Basin bristlecone pines (Pinus longaeva) of California's White Mountains with tree ring counts of over 4,000. A specimen of the magnificent alerce tree (Fitzroya cupressoides) in Chile has recently been shown to have ring counts of over 3,600 years, and some Saharan cypress (Cupressus dupreziana) in Algeria are probably over 2,500 years old. These species all form rings and are arborescent species, having a single trunk. Interesting also, they are all conifers. Tropical trees don't form dependable dating rings, so despite the great size and age that some of these attain, they are bystanders in the contest. Also excluded are those species that aren't trees. Clump forming shrubs or herbaceous perennials, such as azaleas and iris, can persist for many generations and slowly increase their size and numMICHAEl DoSMANN ber of stems. The limits of age on shrubs are unknown, although some documented plantings in botanic gardens are well into their second century. In fact, some of the oldest plants may appear as entire forests or large assemblages of individuals. Many poplar (Populus) species, including quaking aspen (P. tremuloides), can send up multiple individual trunks from a single vast spreading and interconnected root system. These colonies can expand and contract over time depending upon competition, climate, and catastrophic events such as forest fires. Entire mountainsides have been revealed to be covered by a single clonal stand of many trunks connected by a subterranean network of roots. With clonal colonies such as these, estimating age is next to impossible, though in two notable cases this has been attempted. one is the box huckleberry, as described in this article. The other is creosote bush. The shrub Larrea tridentata, known as creosote bush because of its prodigious production of resin, grows in arid regions of the southwestern United States and north central Mexico. Growth of creosote bush colonies begins with the original founding event, the germination of a seedling. As the plant grows, its lower branches come in contact with the soil and develop their own roots. over time the interior portions of the clump die and a ring of plants, slowly increasing in diameter over time, is formed. In the 1980s, botanist Frank Vasek radiocarbon dated chunks of deadwood at the centers of the oldest and largest rings and derived an average growth rate for creosote bush in his region. By applying this rate to the largest clone (for which he found no wood at its epicenter) a phenomenal figure of 9,400 years was obtained (Vasek 1980). This champion plant is now Annual growth rings can be seen in the stems of woody plants in temknown as \"King Clone\" and is properate or arctic regions. A cut trunk of common alder (Alnus glutinosa) is seen here. tected on a 17-acre preserve. Box Huckleberry 15 In 1919, the botanist Frederick Coville postulated (on the basis of morphological characteristics and the inability of the plant to set viable seed) that the large 1,200-foot-long (366 meters) stand in Pennsylvania seemed to be a single clone and had spread across the gentle slope by means of underground runners. It was observed that the plant grew laterally about 6 inches (15.2 centimeters) a year so he estimated that the entire clump had incrementally increased to its present size from a single seed deposited 1,200 years prior. A sister clump, across the Juniata River from the Baird stand, was found in 1920 by H. A. Ward. This was the largest single stand ever to be found, a massive colony stretching over a mile and covering 100 acres (40.5 hectares). Coville's methodology was applied to this monster and an age estimate of 13,000 years was declared. Based on fossil pollen studies, we now have a clearer picture of what the climate and flora of this area would have been like over 13,000 USDA-NRCS PlANTS DATABASE Box huckleberry covers the forest floor at this Pennsylvania site. (photo by rob nicholson) Botanical illustration of Gaylussacia brachycera from Britton and Brown's An illustrated flora of the northern United States, Canada, and the British Possessions, published in 1913. years ago, and these data alone would probably debunk the age claim. The leading edge of the glacier terminated about 75 miles (120.7 kilometers) to the north of the position of the goliath clump around 18,000 to 20,000 years before the present day. As little as 10,000 years ago central Pennsylvania was covered in a boreal forest association, one that would probably have been too cold for the box huckleberry. The current forest, a mix of conifers and deciduous species, started to come into place about 8,000 years ago. In the years since Coville's conjecture, the interstate highway system has had more impact on the plant than any glaciers. During the 1960s a large portion of the goliath colony was eradicated by the installation of US Route 22\/322, and a forest fire also diminished it. Sadly, this construction predated the stronger environmental standards in place today in Pennsylvania, which require highway contractors to inquire about rare and endangered plants in their paths. The conservation status of box huckleberry varies among the states where it is found, but in Pennsylvania it has a ranking of S1--critically imperiled. The species global conservation status, which considers all populations in total, is G3--vulnerable. The tract of land where the original Baird clump grows is now a Pennsylvania State Park while the remnants of the goliath clump are in private hands. The owner is aware of the plant's legacy and seems proud to direct the botanically inclined to its location. 16 Arnoldia 68\/3 CoURTESy oF PlANTENTUIN ESVElD WWW.ESVElD.Nl Box huckleberry's evergreen leaves and delicate bell-shaped flowers are highly ornamental. A Short Walk Through a Short Giant I contacted him, got precise directions, and made a weekend pilgrimage to collect research material from this diminished Methuselah. The two Pennsylvania stands of box huckleberry are in the upper end of the ridge and valley system that stretches from Pennsylvania to Alabama. A drive along the interstate brought me to the top of a bedrock fold overlooking the broad and muddy Juniata River. As directed, I turned from the river and walked into the woods. It was a plain piece of land, a common mix with red maple (Acer rubrum), hickory (Carya spp.), chestnut oak (Quercus prinus), white pine (Pinus strobus), and eastern hemlock (Tsuga canadensis) standing tallest, while below these grew shadblow (Amelanchier spp.) with highbush and lowbush blueberries (Vaccinium corymbosum, V.angustifolium). It would have been quite possible, if you weren't keyed onto it, to walk past the box huckleberry thinking it a variant of lowbush blueberry, or not noticing it at all. Its thick, bright green, leathery leaves are held on wiry pinkish-green stems, and the small, bell-shaped, white and pink flowers are borne in clusters. Its fruit could easily pass for a blueberry, at least until they are chewed, at which point their larger and coarser seeds interrupt the anticipated gastronomic explosion of blueberry deliciousness. In the filtered shade the box huckleberry grew to a height of 8 inches (20.3 centimeters) and formed a patchy patch, denser in some sections than in others. The outer edge of the colony was amoeba-like, its edge curving in and out through the trees. Nearest to the highway was an area of woods that had recently burned, and here the box huckleberry had leaves of a more anemic green, perhaps sun-scorched from want of a shady canopy. After some concentrated tramping, I found another small patch on the slope of a neighboring ridge and sampled this also over two transects. Along with the Baird stand across the river, this would make a total of three separate Pennsylvania populations in the study. I had questioned whether genetic analysis could tease apart these stands to determine whether these large clumps were indeed a single individual run amok or were many indi- Box Huckleberry 17 RoB NICHolSoN Box huckleberry forms a sprawling carpet of green in shaded woodlands. viduals that had merely coalesced together. Dr. Margaret Pooler of the United States National Arboretum had begun some genetic analysis of the species and we agreed that this clonal analysis would make an interesting research subject and also help in determining conservation strategies for this rare species. I established two perpendicular transects across the length and breadth of the clump and then sampled at equal distances along these. I brought cuttings back to the Smith College Botanic Garden, rooted them, and kept them all in separate pots as they grew. The analysis of each sample's genes would show how closely related each of the samples was to one another. If identical in genetic makeup then Coville's single-clone theory would gain credence. To complete the study, material was also collected from a stand in north central Tennessee. The lovely town of Rugby is a quaint cluster of 20 Victorian homes in the woods, a former utopian community now under siege by antique hounds. A path through the woods leads to the Gentlemen's Swimming Hole, and here, growing with the spectacular mountain stewartia (Stewartia ovata), is another outsized patch of box huckleberry, which was also transected and collected for the study. Finally, a sample of the low-growing Mexican Vaccinium kunthianum was also sent to the United States National Arboretum for analysis. Little Plant, Big Data After the team at the National Arboretum did their genetic analysis they found the Tennessee stand was a single clone and the Mexican material was very distantly related. The Baird stand of Pennsylvania showed only two clones, with one clone limited to one corner of the huge clump. Those from across the river were very different. The smaller of the two showed three closely related clones. But a quarter mile away, the largest clump of all--at nearly 1,000 feet (over 300 meters)--showed but one clone. Using the estimated growth rate of 6 inches (15.2 centimeters) per year, this would make this stand 1,000 years old had it started in the middle but 2,000 years old had it begun at the end. Because of the destruction of 80% of the stand we will never know if the entire mile 18 Arnoldia 68\/3 RoB NICHolSoN Coville, F. 1919. The threatened extinction of box huckleberry. Science 50(1280): 3034. Floyd, J. 2002. Phylogenetic and biogeographic patterns in Gaylussacia (Ericaceae) based on morphological, nuclear DNA, and chloroplast DNA variation. Systematic Botany 27 (1): 99115. Gustafson, S. 1986. The Methuselah bush. Rangelands 8 (4): 168169. Kron, K., E. Powell, and J. luteyn. 2002. Phylogenetic relationships within the blueberry tribe (Vaccinieae, Ericaceae) based on sequence data from MATK and nuclear ribosomal ITS regions, with comments on the placement of Satyria. American Journal of Botany 89: 327336. Moldenke, H. 1950. The oldest flowering plant. Horticulture 18: 436437. Pooler, M., l. Dix, and R. Griesbach. 2006. Genetic diversity of the endangered box huckleberry (Gaylussacia brachycera) based on AFlP markers. Journal of the Torrey Botanical Society 133: 3439448 Rooted cutting of Gaylussacia brachycera. Pooler, M., R. Nicholson, and A. Vandegrift. 2008. Clonal fidelity in large colonies of Gaylussacia brachycera Gray (box huckleberry) assessed by DNA fingerprinting. Northeastern Naturalist 15(1): 6774. Sleumer, H. 1967. Die gattung Gaylussacia. Botanische Jahbucher 86: 309384. Smith, H. and D. Smith. 1971. The box huckleberry, Gaylussacia brachycera. Castanea 36: 8189. Vasek, Frank C. 1980. Creosote bush: long-lived clones in the Mojave Desert. American Journal of Botany. 67(2): 246255. Ward, H. 1920. A new station for Gaylussacia brachycera. Rhodora 22: 167168. Wherry, E. 1934. The box huckleberry as an illustration of the need for field work. Bulletin of the Torrey Club 61:8184. Wherry, E. 1972. Box huckleberry as the oldest living protoplasm. Castanea 37: 9495. Wilbur, R.l. and S. Bloodworth. 2004. Notes on the box huckleberry, Gaylussacia brachycera (Ericaceae), and its unexpected presence in North Carolina. Rhodora 106: 371377. Willaman, J. 1965. The oldest living plants. Morris Arboretum Bulletin 16: 6566. Rob Nicholson is Conservatory Manager at the Smith College Botanic Garden in Northampton, Massachusetts. He was formerly Assistant Plant Propagator at the Arnold Arboretum and collected from multiple populations of Gaylussacia brachycera for the Arboretum's collection. length was once all connected and genetically identical. But what was left of Coville's \"charming little thousand-year-old lady of the forest\" may indeed be the oldest known woody plant east of the Rocky Mountains. Space and time are key concerns of our human species and we tend to measure other species by our own familiar rulers and clocks. These large clonal plant stands put the lie to the idea that plants do not move or locomote. A plant specimen will cover distance, but in an imperceptible fashion relative to our lifespan and our ways of moving. It just moves to a slower, millennial-scale timepiece rather than the sweeping second hands that so many of us caffeine-addled commuters adhere to. Perhaps we should all aspire to so relaxed a pace. Bibliography Camp, W.H. 1941. Studies in the Ericales: a review of the North American Gaylussacieae: with remarks on the origin and migration of the group. Bulletin of the Torrey Club 68: 531551. Claypole, E.W. 1883. Note on a Relic of the Native Flora of Pennsylvania, Surviving in Perry County. Proceedings of the American Philosophical Society. 21(114): 226230. "},{"has_event_date":0,"type":"arnoldia","title":"General Forest Ecological Processes chapter 10 from Trees and Forests, a Color Guide","article_sequence":3,"start_page":19,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25506","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070bb26.jpg","volume":68,"issue_number":3,"year":2011,"series":null,"season":null,"authors":"Thomas, Peter A.","article_content":"General Forest Ecological Processes chapter 10 from Trees and Forests, a Color Guide Peter A. Thomas IntroductIon W ooded land currently covers between 30 and 35% of the world's land surface (depending on what is counted as forest), or around 39 to 45 million k2 [15 to 17 million square miles] (FAO 2003). Ecologists often distinguish between woodland and forest. Woodland is a small area of trees with an open canopy (usually defined as the canopy giving less than 40% cover, that is 60% or more of the sky is visible) so that plenty of light reaches the ground, encouraging other vegetation beneath the trees. By contrast, a forest is usually considered to be a relatively large area of trees forming a closed, dense canopy. For simplicity's sake, and because the underlying ecological processes at work are the same, in this chapter the term forest will be used to mean any wooded land. All phOtOs By thE AuthOr A dense temperate rain forest in the Olympic Peninsula, Washington State, USA. It is composed mostly of Psuedotsuga menziesii (Douglas fir), Tsuga heterophylla (western hemlock), and Thuja plicata (western red cedar). 20 Arnoldia 68\/3 SIzE and Growth the most obvious factor that separates forests from other types of habitat is the large weight or mass of organic material present, referred to as the biomass (or sometimes the standing crop). In most forests, more than 85% of the biomass is contained in the above-ground portion of the woody plants. Biomass above ground increases from the northern boreal forest southwards towards the tropics, starting from very low levels at the Arctic tree Open woodland at Needwood Forest, England. The sparse canopy of Fraxinus excelsior (European ash) and Tilia spp. (lime [linden]) line, and reaching in excess of 940 standards allows abundant light to reach the shrub layer of cop- t ha -1 [838,480 pounds per acre] piced Corylus avellana (hazel) and, on the ground, a mixed field in the Amazon basin. however, layer dominated by Hyacinthoides non-scripta (bluebell). A there are exceptionally large forsparse ground layer of mosses is also present. ests outside the tropics, notably the temperate forests of the pacific Northwest of North America. these include stands of huge Psuedotsuga menziesii (Douglas fir), reaching 1,600 t ha-1 [1,427,200 pounds per acre], and Sequoia sempervirens (coastal redwoods), the tallest trees in the world, which have a biomass of up to 3,450 t ha-1 [3,077,400 pounds per acre] just in the trunks. Below-ground biomass in roots is significantly less (Jackson et al. 1996), averaging 29 t ha-1 [25,868 pounds per acre] in boreal forests, 40 to 42 t ha-1 [35,680 to 37,464 pounds per acre] in temperate and tropical deciduous forests, and 49 t ha-1 [43,708 pounds per acre] in tropical evergreen forests. Biomass is a static measure of how much mass there is at any one time, with no indication of how quickly new growth is being added or lost, and so gives little insight into how the forest is functioning. More useful are estimates of the productivity of the forest, i.e. how much new material is being added per year, described as net primary productivity (Npp). this can vary from as little as 1 t ha-1 y-1 [892 pounds per acre per year] in cold boreal forests, to over 30 t ha-1 y-1 [26,760 pounds per acre per year] in tropical rainforests, with an average of 7 to12 t ha-1 y-1 [6,244 to 10,704 pounds per acre per year] in temperate forests. however, a maximum of 36.2 t ha-1 y-1 [32,290 pounds per acre per year] has been recorded in the pacific Northwest from a 26-year-old forest of Tsuga heterophylla (western hemlock). these figures have sometimes been used to calculate how much additional forest needs to be planted to soak up (sequester) the huge amount of extra carbon that is being pumped into the atmosphere--usually approximately 25% extra forest globally. however, such an estimate is blatantly wrong. When a forest is mature it reaches an approximately steady state of mass, where Npp is balanced by an equal loss in biomass through decomposition. At this point, the productivity of the whole forest (the net ecosystem productivity - NEp) drops to near zero. thus, it is only young forests that are carbon Trees and Forests, a Color Guide 21 sinks; once forests are mature they become carbon neutral. In reality, temperate and northern forests globally are a net sink of carbon, but this is primarily due to expansion of the amount of forest due to reforestation (Beedlow et al. 2004). LIGht trees have evolved as a life form to outcompete their neighbors for light by growing tall, so producing dense forests that inside are darker, more humid, and less prone to extremes of temperature variation than outside. In temperate forests at least, it is usually possible to recognize four reasonably distinct layers. At the top is the tree canopy, normally 5+ m [16.4 feet or greater] above ground. Below are the shrub layer (< 5 m) [less than 16.4 feet], the field or herb layer of herbaceous plants and short woody plants such as brambles, and the ground or moss layer of mosses and liverworts, lichens, and algae. Each layer blocks sunlight so that a dense layer may preclude any layers below, and the forest floor may be very dark indeed. In temperate regions, the amount of light reaching the forest floor may be as high as 20 to 50% of full sunlight in an open birch wood, down to just 2 to 5% beneath Fagus sylvatica (European beech). In these deciduous forests, light levels are higher once the leaves have fallen, but the trunks and branches still block some light such that light levels are likely to be below 70 to 80% of full sun. Evergreen forests Primula vulgaris (primrose), a wintergreen plant that keeps tend to cast similar shade all year round; some leaves alive throughout the year. in Europe, light levels below natural Pinus sylvestris (scots pine) forests are usually around 11 to 13%, while below Picea abies (Norway spruce) they can be as low as 2 to 3%. In tropical rain forests, light levels at the forest floor may be even lower, just 0.2 to 2% of full sunlight. As a rule of thumb, plants require 20% of full sunlight for maximum photosynthesis and at least 2 to 3% sunlight for photosynthesis to exceed background respiratory costs (the compensation point). this inevitably means that the floor of densest forests is at, or beneath, the limits of plant growth. some forest floor plant specialists have overcome this problem with a number of physiological solutions. using shade leaves that are thinner and more efficient at low light levels than sun leaves. reducing the compensation point. Bates and roeser (1928) found that coastal redwood in deep shade requires just 0.62% sunlight. Making use of sunflecks--patches of sunlight passing through gaps in the canopy--which can briefly give up to 50% of full sunlight and make up 70 to 80% of the total solar energy reaching the ground in a dense forest (Evans 1956). these flecks are especially important to shade plants that are capable of responding quickly to the brief flurries of light. 22 Arnoldia 68\/3 plants can also cope with dark conditions by avoidance. temperate deciduous forests are well-known for their colorful carpets of prevernal plants, which grow and flower early in spring. In the uK these include Hyacinthoides non-scripta (bluebell), Ranunculus ficaria (lesser celandine), and Anemone nemorosa (wood anemone). these plants make use of the light reaching the ground before the trees develop their canopy of leaves, and die back once the shade is too deep. summergreen plants, such as Mercurialis perennis (dog's mercury) and Galium odoratum (woodruff), are similar but keep their leaves through the summer using what little light is available. As an extension of this strategy, wintergreen plants (which keep at least a few green leaves all year round) and true evergreen plants can start growth as soon as spring conditions allow, and continue growth into a warm late autumn after leaf fall. such plants include wintergreen Oxalis acetosella (wood sorrel) and Primula vulgaris (primrose), and evergreens such as Hedera helix (ivy) and Ilex aquifolium (holly). Being evergreen is an efficient strategy for coping with seasonally abundant light, but it does carry costs. In winter, holly is a sitting target for herbivores such as deer, and so has evolved prickly spines to the leaves. these spines are absent above deer-browsing height, around 3 m [9.8 feet] above ground. tree seedlings face similar problems of shade, having to grow up through dark layers of vegetation before reaching the canopy. Different tree species vary tremendously in how much shade they can bear as seedlings and saplings. Fagus sylvatica (European beech) and Acer saccharum (sugar maple, from North America) are very tolerant of deep shade, while Betula spp. (birches) and Populus spp. (poplars) grow best under high light intensities. however, it is now apparent that the ability to tolerate shade can change through the lifespan of a tree (poorter et al. 2005), so it is possible that many trees are more shade tolerant as seedlings than as adults. Nevertheless, comparatively few trees can tolerate the full shade cast by their mature relatives. Consequently, they depend Seedlings of Fagus sylvatica (European beech) are very shade-tolerant and upon gaps appearing in the forest, capable of growing under the dense canopy of their parents. Each seedling by one or more trees dying or fallshows two distinctively shaped cotyledons below the young shoot. ing, for successful establishment of seedlings. Gaps are sufficiently important that while large-scale regional vegetation (e.g., oak forest) is determined by climate, soil, and topography, it is the dynamics of gaps that largely controls the proportions in which the various species grow in any one area. For example, in small gaps created by one tree falling, shade tolerant trees such as Fagus spp. (beech) or Abies spp. (fir) are more likely to do best and dominate. In larger gaps, species such as Betula (birch) and Salix (willow), Trees and Forests, a Color Guide 23 which invade quickly from light, wind-borne seeds and grow rapidly, are more likely to dominate initially but later give way to shade-tolerant trees. It is not just what goes on above ground that is important; in larger gaps there will also be less belowground competition from the root systems of the large trees at the gap edge. the importance of such competition has been demonstrated experimentally by cutting roots (trenching) around the edges of a plot: seedlings inside the plot usually grow faster (e.g., Barberis and tanner 2005). Competition may also happen below ground from the field layer vegetation by allelopathy, i.e., secretion of chemicals, which inhibit other root growth, into the soil (e.g., Orr et al. 2005). Further variability in seedling establishment is produced by small-scale heterogeneity of the forest floor. pits and mounds of bare mineral soil created by falling trees offer less competition and a more constant water supply than the surrounding humus-rich forest floor. In a Pinus sylvestris (scots pine) forest in Finland, Kuuluvainen and Juntunen (1998) found that although these bare sites covered just 8.4% of the forest, they held 60% of pine and 91% of birch seedlings and saplings. Dense field and ground layers can cause problems for tree regeneration, swamping small seedlings. this is one reason why, in temperate rainforests, seedlings are often most common on \"nurse logs,\" which are continuously damp enough to provide moisture and lift the seedlings above the dense field layer. As tree seedlings grow upwards into a gap, there can be intense competition to reach and keep the light; whichever seedlings grow quickest will dominate the gap, at least in the short term. A common strategy to get a head start, found in trees as diverse as Fraxinus excelsior (European ash), and shade-tolerant firs (Narukawa and yamamoto 2001), is to have a seedling bank. here, young plants survive in light conditions below their compensation point (i.e. they are sustaining a net loss of energy) and grow very slowly while their energy reserves last. these seedlings are then able to take rapid advantage of an opening in the canopy in the race for dominance. watEr Given that a single, large deciduous tree can use 400,000 liters [105,670 gallons] of water in transpiration in a summer (thomas 2000), it is obvious that whole forests move immense amounts of water from the soil to the atmosphere. Nevertheless, water is rarely limiting for tree growth in temperate regions until rainfall decreases to such an extent that scrub and grasslands take over. Almost all roots tend to be quite shallow, so potential problems exist if the surface layers of the soil are drained of available water between rain events. this is obviated, however, by the process of hydraulic lifting present in a number of trees and a few grasses. here, water is raised at night from moist areas lower in the soil (flowing along a hydraulic gradient through the roots) to nearer the surface. hydraulic lifting is most common in savannas and other xeric (dry) woodlands, especially among older trees (Domec et al. 2004), but is found elsewhere. the amounts moved can be significant: a mature Acer saccharum (sugar maple) 19 m [62.3 feet] high can raise around 100 liters [26.4 gallons] of water each night compared to a water loss via transpiration of 400 to 475 liters [106 to 125 gallons] the following day (Emerman and Dawson 1996). this raised water also benefits other surrounding plants (penuelas and Filella 2003; Filella and penuelas 20032004). 24 Arnoldia 68\/3 Forests also play a significant role in the redistribution of water on a regional scale. rainfall intercepted by the canopy is evaporated before it reaches the ground. When this and the transpiration of water are combined (evapotranspiration), the overall losses are in the order of 30 to 60% of precipitation in deciduous forests, 50 to 60% in tropical evergreen forests, and 60 to 70% in coniferous forests, compared to around 20% in grasslands. Not surprisingly, forested areas have water yields (measured as stream flow) 25 to 80% lower than pastures. Moreover, computer modeling by Calder et al. (2003) suggests that planting oak woodland in central England would eventually reduce recharge of aquifers and runoff to streams by almost one half. so, should forest be removed to improve water yield? Most data show that regardless of forest type, removal of up to 20% of the trees has an insignificant effect on water yield, presumably because of increased soil evaporation replacing evapotranspiration (Brown et al. 2005). Further clearance does improve water yield (Bosch and hewlett 1982), but by comparatively small amounts until clearance is significant. Many people have held the view that forests increase rainfall in a watershed through evaporating water, thus helping build clouds. however, in temperate areas, at least, the contribution of a forest to rainfall is likely to be insignificant and certainly less than 5% (Golding 1970). On a continental scale, forests help to increase rainfall in the sense that they repeatedly recycle the atmospheric moisture passing from the oceans to the land. For example, in the Amazon Basin, much of the daily rainfall is immediately evaporated to generate clouds for rainfall downwind. It is highly likely that continual clearance of the forest will reduce rainfall elsewhere in the region since much of the water will enter rivers and be lost to the system. Moreover, the effects of such tropical deforestation have far wider repercussions in mid- and high latitudes through large-scale links in the water cycle and weather. Avissar and Werth (2005) have shown, for example, that deforestation of Amazonia and Central Africa severely reduces rainfall in the Midwest of the united states. nutrIEntS Nitrogen is usually the nutrient most limiting growth in temperate forests, while in other forests, especially on soils of great age, phosphorus may well be the limiting nutrient. Nutrients within a forest ecosystem are highly recycled and key to this recycling are the decomposer organisms that release nutrients from dead material. larger soil fauna, such as earthworms and beetles, chew debris into fine particles suitable for the soil fungi and bacteria. A square meter of soil in temperate woodland may contain more than 1,000 species of animal, from protozoa to earthworms, and a gram of soil can contain more than 1,000 species and more than 200 million bacterial cells (Fitter, 2005). soil organic matter (surface litter and humus incorporated into the soil) is thus the main bottleneck controlling nutrient availability to plants, and the slower decomposition is, the more of a limiting factor it is. this helps explain why slow plant growth occurs on cold northern soils that have large organic matter accumulations. Fungi and bacteria are not altruistic in providing nutrients to plants. As dead material is decomposed, nutrients released by the microorganisms are immediately taken back up by other microorganisms, and so are effectively immobilized and unavailable to plants. however, as the carbon is progressively used up in their Trees and Forests, a Color Guide 25 respiration (and released as carbon dioxide), the conserved nutrients become more than the microbes can use, and the excess is released in inorganic form for plants to use. Consequently, when a fresh batch of litter arrives on the forest floor there is a variable time lag before its carbon has been reduced sufficiently to allow nutrients to be freed into the soil for plant growth, the process being regulated by the microbial community (Attiwill and Adams 1993; Agren et al. 2001). plants can, however, circumvent this bottleneck in several ways. Firstly, more than 80% of the world's vascular plants have on their roots mycorrhizal fungi, which greatly assist in scavenging nutrients from the soil to the symbiotic benefit of both plants and fungi. secondly, some plants are now known to be able to directly use organic nutrients, without the intervention of microorganisms first breaking them down into inorganic forms. For example, up to 50% of the total nitrogen in forest soils is usually in the form of dissolved organic nitrogen (DON), of which approximately 10 to 20% consists of amino acids. the degree to which plants can use DON is open to speculation, but it is becoming clear that many plants are capable of absorbing amino acids directly (lipson and Nasholm 2001) and are thus able to short-circuit the microorganism bottleneck. the same may also be true for organic phosphorus. Although nutrients are tightly recycled within a forest ecosystem, there are still (usually small) annual inputs and losses. Nutrients are added to forests The effect of 14 years of nitrogen enrichment on Pinus resinosa (red pine) at Harvard Forest: (top) control plot with no extra nitrogen added above the background deposition of 7 to 8 kg N ha-1 y-1; (middle) low N addition (50 kg N ha-1 y-1); and (bottom) high N addition (150 kg N ha-1 y-1). 26 Arnoldia 68\/3 through rain and dust, dissolved from rocks in the soil, and as biological input from nitrogen fixation by microbes. losses of nutrients can be very rapid due to fire, wind, and erosion but the majority of losses, from temperate forests at least, are by leaching of nutrients as water percolates through the soil. however, since nutrients are vital to forest growth, plants and microbes are fairly efficient at reabsorbing and holding available nutrients and creating conditions of controlled decomposition. this has been admirably demonstrated by the hubbard Brook Ecosystem study in the White Mountain National Forest of New hampshire, established in 1963 (likens 2004). As part of this, a discrete watershed was clear-felled in 19651966 and treated with herbicides for three years to prevent any regrowth, while a similar watershed had the hardwood forest left intact. After clear-felling, stream flow went up (due to reduced evapotranspiration) and net losses of nitrate, calcium, and potassium in stream water generally peaked in the second year, each returning to pre-cutting levels at rates unique to each ion as the forest regrew. however, even decades after clear-felling, differences in stream water solutes can still be seen, especially in calcium (likens et al. 1998). there is still a good deal to learn about mechanisms of nutrient retention in forests. For example, Muller and Bormann put forward the vernal dam hypothesis in 1976. this proposes that prevernal plants, which grow early in spring before canopy closure, take up nitrogen and other nutrients before they can be leached; these are subsequently made available to other plants as the prevernal plants die back from lack of light. At hubbard Brook, plants of Erythronium americanum (yellow trout lily) saved almost half of the important nutrients from being washed away. In the spring they used 43 and 48% of the released potassium and nitrogen, respectively, with the rest being lost in stream water. some subsequent experiments (e.g., tessier and raynal 2003) have supported the theory. however, other contradictory studies have shown that the microbe population itself is better at soaking up the spring burst of nutrients (e.g., Zak et al. 1990). Also, while the dying back of vernal plants can produce a burst of nutrients (e.g., Anderson and Eickmeier 2000), the plants may not be very efficient at taking up nutrients in the first place (e.g., Anderson and Eickmeier 1998; rothstein 2000). undoubtedly, some of the experimental differences come from investigating different plant species in several forests. the tight recycling of nutrients within the forest ecosystem can cause problems if too much arrives as pollution. Nitrogen enrichment, particularly in northern temperate areas, is just such a case (Nosengo 2003). since the 1980s, normal background nitrogen deposition of < 1 kg ha-1 y-1 [less than .892 pounds per acre per year] has increased by 10 to 40 times or even higher. the effect of too much nitrogen is clearly seen in long-term experiments running at harvard Forest, Massachusetts since 1988 (Magill et al. 2004). In one of these, a plantation of Pinus resinosa (red pine) was subjected to three levels of nitrogen: a control, low N addition, and high N addition. After 14 years, annual wood production had decreased by 31% and 54% relative to the control in the low N and high N plots, respectively, and the canopies had thinned due to dieback under higher nitrogen levels. Mortality also increased (control 12%; low N 23%; high N 56%) and the whole high N stand was expected to die in the near future. Trees and Forests, a Color Guide 27 coarSE woody dEbrIS the vital importance of dead wood in forest carbon budgets, and also as an invaluable wildlife resource, has been increasingly appreciated over the last decade (Kirby and Drake 1993). Dead wood appears in many forms, sizes, and positions including standing dead trees (snags), dead branches in the canopy, and trunks and branches on the ground. A useful term for this motley collection is coarse woody debris (CWD). typically, CWD in a forest forms up to a quarter of all the above-ground biomass and is normally in the range of 11 to 38 t ha-1 [9,812 to 33,896 pounds per acre] in deciduous forests, with the largest amounts in cooler regions where decomposition is slower. Conifer forests generally hold more CWD than deciduous forests, typically around 100 t ha-1, [89,200 pounds per acre] but up to 500 t ha-1 [446,000 pounds per acre] in the coastal redwood forests of California and the rain forests of the pacific Northwest. tropical forests, with more rapid decomposition, usually have lower amounts of woody accumulation, but levels up to 100 t ha-1 [89,200 pounds per acre] are possible in more water-logged areas of the Amazonian forest. If 100 t ha-1 [89,200 pounds per acre] of wood was spread evenly over the forest floor it would amount to 10 kg [22 pounds] in each square meter. however, because the bulk of the wood is in large pieces, typically less than 5% of the ground will be covered by CWD, although this can rise to around a third cover in very dense coniferous forests. snags are of particular wildlife interest. In the Bialowiea forest of poland, one of the most pristine forests in Europe, Bobiec (2002) found that standing dead wood varied from 3 to 21% of total CWD, and figures of 25% are typical in many of the world's forests. Wood is difficult to decompose. It is composed of 40 to 55% cellulose, 25 to 40% hemicelluloses, and 18 to 35% lignin (conifers having a greater proportion of lignin than hardwoods). Wood is thus high in structural carbohydrates (which require specialized enzymes to break them up) but also poor in nutrients such as nitrogen: 0.03 to 0.1% N (by mass) compared to 1 to 5% in foliage. In most forests, wood (CWD) will be colonized by fungi within a year and completely colonized within 5 to 10 years. however, decay rates of wood vary tremendously depending upon the climate, decaying organisms available, and the size and type of wood. In general terms, pioneer trees such as birches and willows Temperate rain forests, such as the one here on western Vancouver Island, Canada, can contain large quantities of dead wood, in part invest less energy in protecting because of the size of some of the fallen logs. The one shown here is of their wood from rot (going for Picea sitchensis (Sitka spruce). 28 Arnoldia 68\/3 speed of growth rather than defense) and logs on the ground rot away within a few decades. Wood from longer-lived trees such as oaks may persist for a century or much longer, while in cool climates such as the pacific Northwest wood may persist for up to 600 years (Franklin et al. 1981). Even in tropical rain forests, wood above 3 cm [1.2 inches] diameter takes at least 15 years to decompose (Anderson and swift 1983). Again, however, environmental conditions play an important role in determining decay rates; logs of Populus balsamifera (balsam poplar) in North America, which would decay away within 40 to 60 years on land, last for over 250 years when waterlogged in a beaver pond. EvErGrEEn and dEcIduouS LEavES At first sight, the occurrence of evergreen and deciduous trees in different forests can appear haphazard, but in reality it demonstrates the interactions of many of the ecological processes described above (thomas 2000). Deciduous trees lose their leaves during an unfavorable season (winter in temperate areas), while evergreen trees always have some leaves on the tree and individual leaves Evergreen conifers, such as Abies lasiocarpa (submay live from six months to over 30 years. If grow- alpine fir) shown here in the Canadian Rocky Mountains, are typical of areas with short growing seasons ing conditions are favorable all year round, as in where deciduous trees are disadvantaged by wasting tropical rain forests, then there is no selective part of the season producing new leaves. advantage in being deciduous and so evergreen angiosperms dominate. In climates with a dry summer or cold winter, it is cheaper to grow thin disposable leaves than to grow more robust leaves capable of surviving the offseason, so in most moist temperate areas deciduous trees dominate. however, if environmental conditions become worse, it may once again be more beneficial to grow evergreen leaves. this includes areas with a very short growing season, where evergreen leaves are able to start growing as soon Deciduous forest in Harvard Forest, Massachusetts, USA. In a seasonal temperate climate it is more economical for trees to grow a set of disas conditions allow and so none posable leaves each spring rather than build leaves capable of surviving of the growing season is wasted. the winter. Trees and Forests, a Color Guide 29 this accounts for evergreen leaves in northern and alpine areas, and also among woodland understory shrubs such as holly and ivy, which benefit from an early spring start and late autumn finish when the canopy has no leaves. Evergreen leaves are also found in Mediterranean climates where the winter growing season is dry; leaves that are protected enough to cope with the droughty conditions will also sur- In the very short growing season of the tundra overlying permafrost, evergreen shrubs give way to deciduous Salix spp. (willows). (The graves are those of 19th century vive the hot dry sum- whalers who overwintered and died here on Herschel Island in the Arctic Ocean.) mer, and so effectively become evergreen and need to be kept for several years to repay the high investment cost. In areas where the climate becomes even more severe, such as at the Arctic tree line or in alpine areas, deciduous leaves re-appear. Despite the problems of a very short growing season and acute shortage of nutrients, the winter is so severe that it is cheaper to build new leaves every year rather than attempting to keep leaves alive. thus, the northernmost trees in the Arctic and uppermost trees in alpine areas are deciduous trees such as species of Betula (birch), Larix (larch), and Salix (willow). concLudInG rEmarkS Forest ecosystems work in much the same way as any other ecosystem, but size and complexity create ecological situations that are unique to forests. the large amounts of biomass that can be grown in a year appear useful for carbon sequestration in relation to global warming, but must be weighed against the decompositional losses in mature forests, and possibly the extra methane--a potent greenhouse gas--that these will generate (Keppler et al. 2006). to maintain sequestration rates, new forests are constantly needed. light availability presents problems for those plants living below the dense forest canopy, but these problems are solved by making do with less light or growing when light is available in the spring or during brief sunflecks. the role of forests in the water cycle still needs to be fully clarified, but it is of great importance due to the likely pressure on forests as human water needs increase. Nutrient dynamics in forests are crucial to their long-term well-being and it is important that we improve our understanding of the effects of climate change and pollution on decomposition and nutrient cycling. Of necessity, this chapter gives only a resume of a very large subject. A more detailed account of forest ecology is provided by thomas and packham (2007). 30 Arnoldia 68\/3 rEFErEncES Agren, G. I., E. Bosatta, and A.h. Magill. 2001. Combining theory and experiment to understand effects of inorganic nitrogen on litter decomposition. Oecologia 128: 9498. Anderson, J.M. and M.J. swift. 1983. Decomposition in tropical forests. In: sutton, s.l., t.C. Whitmore, and A.C. Chadwick (eds.), Tropical Rain Forest: Ecology and Management. Blackwell scientific publications, Oxford, pp.287309. Anderson, W.B. and W.G. Eickmeier. 1998. physiological and morphological responses to shade and nutrient additions of Claytonia virginica (portulacaceae): implications for the \"vernal dam\" hypothesis. Canadian Journal of Botany 76: 13401349. Anderson, W.B. and W.G. Eickmeier. 2000. Nutrient resorption in Claytonia virginica l.: implications for deciduous forest nutrient cycling. Canadian Journal of Botany 78: 832 839. Attiwill, p.M. and M.A. Adams. 1993. Nutrient cycling in forests. New Phytologist 124: 561582. Avissar, r. and D. Werth. 2005. Global hydroclimatological teleconnections resulting from tropical deforestation. Journal of Hydrometeorology 6: 134145. Barberis, I.M. and E.V.J. tanner. 2005. Gaps and root trenching increase tree seedling growth in panamanian semi-evergreen forest. Ecology 86: 667674. Bates, C.G. and J. roeser Jr. (1928). light intensities required for growth of coniferous seedlings. American Journal of Botany, 15: 185194. Beedlow, p.A., D.t. tingey, D.l. phillips, W.E. hogsett, and D.M. Olszyk. 2004. rising atmospheric CO2 and carbon sequestration in forests. Frontiers in Ecology and the Environment 2: 315322. Bobiec, A. 2002. living stands and dead wood in the Bialowiea forest: suggestions for restoration management. Forest Ecology and Management 165: 125140. Bosch, J.M. and J.D. hewlett. 1982. A review of catchment experiments to determine the effect of vegetation changes on water yield and evapotranspiration. Journal of Hydrology 55: 323. Brown, A.E., l. Zhang, t.A. McMahon, A.W. Western, and r.A. Vertessy. 2005. A review of paired catchment studies for determining changes in water yield resulting from alterations in vegetation. Journal of Hydrology 310: 2861. Calder, I.r., I. reid, t.r. Nisbet, and J.C. Green. 2003. Impact of lowland forests in England on water resources: application of the hydrological land use Change (hyluC) model. Water Resources Research 39: paper 1319. Domec, C., J.M. Warren, F.C. Meinzer, J.r. Brooks, and r. Coulombe. 2004. Native root xylem embolism and stomatal closure in stands of Douglas-fir and ponderosa pine: mitigation by hydraulic redistribution. Oecologia 141: 716 Emerman, s.h. and t.E. Dawson. 1996. hydraulic lift and its influence on the water content of the rhizosphere: an example from sugar maple, Acer saccharum. Oecologia 108: 273278. Evans, G.C. 1956. An area survey method of investigating the distribution of light intensity in woodlands, with particular reference to sun flecks, including an analysis of data from rain forest in southern Nigeria. Journal of Ecology 44: 391428. FAO. 2003. State of the World's Forests 2003, Food and Agriculture Organization of the united Nations, rome, Italy. Filella, I. and J. penuelas. 20032004. Indications of hydraulic lift by Pinus halepensis and its effects on the water relations of neighbor shrubs. Biologia Plantarum 47: 209214. Fitter, A.h. 2005. Darkness visible: reflections on underground ecology. Journal of Ecology 93: 231243. Franklin, J.F., K. Cromack, Jr., A. McKee, C. Masser, J. sedell, F. swanson, and G. Juday. 1981. Ecological Characteristics of Old-growth Douglas-fir Forests. united states Department of Agriculture, Forest service, pacific Northwest Forest and range Experiment station, portland, Oregon, General technical report pNW-118. Golding, D.l. 1970. the effects of forests on precipitation. Forestry Chronicle 46: 397 402. Jackson, r.B., J. Canadell, J.r. Ehleringer, h.A. Mooney, O.E. sala, and E.D. schulze. 1996. A global analysis of root distributions for terrestrial biomes. Oecologia 108: 389411. Keppler, F., J.t.G. hamilton, M. Bra, and t. rockmann. 2006. Methane emissions from terrestrial plants under aerobic conditions. Nature 439: 187191. Kirby, K. and M. Drake. 1993. Dead wood matters. English Nature Science No. 7, English Nature, peterborough. Kuuluvainen, t. and p. Juntunen. 1998. seedling establishment in relation to microhabitat variation in a windthrow gap in a boreal Pinus sylvestris forest. Journal of Vegetation Science 9: 551562. likens, G.E. 2004. some perspectives on long-term biogeochemical research from the hubbard Brook Ecosystem study. Ecology 85: 23552362. likens, G.E., C.t. Driscoll, D.C. Buso, t.G. siccama, C. Johnson, G.M. lovett, t.J. Fahey, W.A. reiners, D.F. ryan, C.W. Martin, and s.W. Bailey. 1998. the biogeochemistry of calcium at hubbard Brook. Biogeochemistry 41: 89173. lipson, D. and t. Nasholm. 2001. the unexpected versatility of plants: organic nitrogen use and availability in terrestrial ecosystems. Oecologia 128: 305316. Magill, A.h., D.A. Aber, W.s. Currie, K.J. Nadelhoffer, M.E. Martin, W.h. McDowell, J.M. Melillo, and p. steudler. 2004. Ecosystem response to Trees and Forests, a Color Guide 31 15 years of chronic nitrogen additions at the harvard Forest ltEr, Massachusetts, usA. Forest Ecology and Management, 196: 728. Muller, r.N. and F.h. Bor mann. 1976. role of Erythronium americanum Ker. in energy flow and nutrient dynamics of a northern hardwood forest ecosystem. Science 193: 11261128. Narukawa, y. and s-I.yamamoto. 2001. Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan. Ecological Research 16: 617625. Nosengo, N. 2003. Fertilized to death. Nature 425: 894 895. Orr, s.p., J.A. rudgers, and K. Clay. 2005. Invasive plants can inhibit native tree seedlings: testing potential allelopathic mechanisms. Plant Ecology 181: 153165. penuelas, J. and I. Filella. 2003. Deuterium labeling of roots provides evidence of deep water access and hydraulic lift by Pinus nigra in a Mediterranean forest of NE spain. Environmental and Experimental Botany 49: 201208. poorter, l., p.A. Zuidema, M. pena-Claros, and r.G.A. Boot. 2005. A monocarpic tree species in a polycarpic world: how can Tachigali vasqueszii maintain itself in a tropical rain forest. Journal of Ecology 93: 268278. rothstein, D.E. 2000. spring ephemeral herbs and nitrogen cycling in a northern hardwood forest: an experimental test of the vernal dam hypothesis. Oecologia 124: 446453. tessier, J.t. and D.J. raynal. 2003. Vernal nitrogen and phosphorus retention by forest understory vegetation and soil microbes. Plant and Soil 256: 443453. thomas, p. 2000. Trees: Their Natural History. Cambridge university press, Cambridge. thomas, p.A. and J.r. packham. 2007. Ecology of Woodlands and Forests. Cambridge university press, Cambridge. Zak, D.r., p.M. Groffman, K.s. pregitzer, s. Christensen, and J.M. tiedje. 1990. the vernal dam: plant microbe competition for nitrogen in northern hardwood forests. Ecology 71: 651656. Excerpted with permission from: Trees and Forests, a Colour Guide (in united Kingdom) Edited by Bryan G. Bowes 2010, Manson publishing ltd. IsBN 978-1-84076-085-9 published in the united states as: Trees and Forests, a Color Guide Academic press\/Elsevier IsBN 978-0-12-382173-7 Note: Not all photographs from the book are included in this excerpt. peter A. thomas is senior lecturer in Botanical and Environmental sciences in the school of life sciences, Keele university. 36673667 U.S. POSTAL SERVICE STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION (Required by 39 U.S.C. 3685) 1. publication title: Arnoldia. 2. publication No: 00042633. 3. Filing Date: september 30, 2010. 4. Issue Frequency: Quarterly. 5. No. of Issues published Annually: 4. 6. Annual subscription price: $20.00 domestic; $25.00 foreign. 7. Complete Mailing Address of Known Office of publication: Arnold Arboretum, 125 Arborway, Boston, suffolk County, MA 021303500. 8. Complete Mailing Address of headquarters of General Business Office of publisher: Arnold Arboretum, 125 Arborway, Boston, suffolk County, MA 021303500. 9. Full Names and Complete Mailing Address of publisher, Editor, and Managing Editor: Arnold Arboretum, 125 Arborway, Boston, suffolk County, MA 021303500, publisher; Nancy rose, Arnold Arboretum, 125 Arborway, Boston, MA 021303500, editor. 10. Owner: the Arnold Arboretum of harvard university, 125 Arborway, Boston, suffolk County, MA 021303500. 11. Known Bondholders, Mortgagees, and Other security holders Owning or holding 1 percent or More of total Amount of Bonds, Mortgages, or Other securities: none. 12. the purpose, function, and nonprofit status of this organization and the exempt status for federal income tax purposes have not changed during the preceding 12 months. 13. publication Name: Arnoldia. 14. Issue Date for Circulation Data Below: June 17, 2010. 15. Extent and Nature of Circulation. a. total No. Copies. Average No. Copies Each Issue During preceding 12 Months: 2,867. Actual No. Copies of single Issue published Nearest to Filing Date: 2,800. b. paid and\/or requested Circulation. (1) paid\/requested Outside-County Mail subscriptions. Average No. Copies Each Issue During preceding 12 Months. Copies Each Issue During preceding 12 Months: 1,430. No. Copies of single Issue published Nearest to Filing Date: 1,428. (2) paid In-County subscriptions. Average No. Copies Each Issue During preceding 12 Months. Copies Each Issue During preceding 12 Months: 396. No. Copies of single Issue published Nearest to Filing Date: 386. (3) sales through Dealers and Carriers, street Vendors, and Counter sales: none. (4) Other Classes Mailed through the usps: none. c. total paid and\/or requested Circulation. Average No. Copies Each Issue During preceding 12 Months: 1,826. Actual No. Copies of single Issue published Nearest to Filing Date: 1,814. d. Free Distribution by Mail. Average No. Copies Each Issue During preceding 12 Months: 108. 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Actual No. Copies of single Issue published Nearest to Filing Date: 75%. I certify that all information furnished on this form is true and complete. Nancy rose, Editor. "},{"has_event_date":0,"type":"arnoldia","title":"A Taste of Sichuan: Zanthoxylum simulans","article_sequence":4,"start_page":32,"end_page":32,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25505","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070b76d.jpg","volume":68,"issue_number":3,"year":2011,"series":null,"season":null,"authors":"Damery, Jonathan","article_content":"A Taste of Sichuan: Zanthoxylum simulans Jonathan Damery S pices have molded the modern world. Columbus and his fleet sailed in the name of black pepper. New Amsterdam, the Dutch colonial settlement on Manhattan Island, became New York in an island trade essentially pertaining to nutmeg. And the records go on back, to the time when the caravan routes of the rising Arab world first introduced exotic Eastern spices to the insipid foods of Europe. The worldwide distribution of spice-producing plant species (and especially the woody plants within that group) is limited mostly to tropical climates, which excludes the most common spice plants from the temperate-zone tree and shrub collections of the Arnold Arboretum. Of the few exceptions, the Arboretum is home to one spice-producing plant of particular interest--Zanthoxylum simulans, a source of Sichuan pepper. Though not a common spice in American kitchens, the signature mouth-numbing flavor of Sichuan pepper is indispensible in the regional cuisine of Sichuan, China. It is produced from the dried fruit of several different species of Zanthoxylum, also known as the prickly ashes. The most common sources are Z. bungeanum and Z. simulans (formerly considered varieties of the same species), and Z. piperitum is used for similar culinary purposes in Japan. IT IS INTERESTINg to note that Zanthoxylum is a member of the citrus family (Rutaceae). In the United States, this relationship has proved to be rather problematic for Sichuan cooks. Thought to be a potential mode of introducing citrus canker to American citrus groves, the United States Food and Drug Administration forbade its importation for nearly forty years, from 1968 to 2005 (though serious enforcement only came about in 2002). The ban was lifted under the stipulation that the Sichuan pepper be heated just enough to kill any infectious bacteria prior to importation. Several examples of both Zanthoxylum simulans (flatspine prickly ash) and Z. piperitum can be found growing in the Arboretum, but the most notable is a large specimen of the former, tucked in just before the smoketree collection on Meadow Road. This specimen (accession 1803-77-A) was collected as seed by Arboretum taxonomists Stephen Spongberg and Richard Weaver in 1977 from the Forest Research Institute in Seoul, South Korea. The gracefully spreading form of this specimen (24.3 feet [7.4 meters] tall, 6.5 inch [16.5cm] diameter main stem) melds easily into the border of Meadow Road, but even in the winter it is worth taking a few steps off the path for a closer view. A spiny plant from twig to trunk, the spines (or technically prickles in the case of those on the trunk) become enlarged and woody, lending an exotic appearance to the tree. The deep green compound leaves have an attractive glossy sheen, and in midsummer Z. simulans is covered with a greenish white haze of small flowers, followed by a prolific display of small, round follicles (a type of dehiscent fruit). At maturity these fruits turn a pinkishbronze color and split open, spitting out the seeds. The dried follicle is the culinary product, Sichuan pepper. On the plains and in upland forests of northern and central China, Arboretum plant explorer E. H. Wilson reported that Zanthoxylum simulans grew naturally on cliffs and waysides. In cultivation it was grown in dry, hot river valleys. This is a good indication of its adaptability to a myriad of difficult landscape conditions. The species is cold hardy in USDA Zones 5 to 7 (average annual minimum temperature -20 to 10F [-28.8 to -12.3C]). From the bare winter trunks through the remarkable autumn fruit display, Zanthoxylum simulans is worth viewing all year long. It is a not-so-hidden--but often missed--Arboretum treasure. Jonathan Damery is a Curatorial Fellow at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23422","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160b728.jpg","title":"2011-68-3","volume":68,"issue_number":3,"year":2011,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Case for Plant Exploration","article_sequence":1,"start_page":2,"end_page":2,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25501","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070af26.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"The Case for Plant Exploration Peter Del Tredici he history of plant exploration is as old as human history itself. People have been discovering, collecting, and moving plants for eons, and the process is not likely to stop any time soon. Indeed, it is as ancient as the practice of agriculture itself--it's part of our genetic heritage. The challenge plant collectors face today is how to continue their work without causing further problems for our already badly damaged environment. Despite the best efforts of many research scientists, we have yet to develop a truly reliable way of predicting whether an unknown plant will be problematic without actually growing it under a variety of conditions to see how it behaves. Botanical gardens, with their relatively secure perimeters and their commitment to science over commerce, are places where new plant introductions can and should be tested for a variety of traits including their potential invasiveness As the world environment continues to deteriorate as a result of human-induced phenomena such as acid rain and climate change, there can be little doubt but that we are going to need tough, adaptable plants for Peter Del Tredici inspecting persimmon seeds our managed landscapes more than ever. Many during the NACPEC expedition to Wudang Shan, of our native species--including such familiar Hubei, China, in 1994. trees as American elm, eastern hemlock, sugar maple, and white and green ash--are no longer planted in our cities because of insect, disease, or stress susceptibility. We have a real need to replace them with stress-tolerant, non-invasive species that can survive all the abuse that people throw at them. Some of these plants of the future may be native to North America, but I can guarantee you that some of them--either as species or as hybrids--will come from Central and Eastern Asia. And that's where the North AmericaChina Plant Exploration Consortium comes in. For the past twenty years this collaborative organization has made it a priority to try to deal with future horticultural problems without creating new ones in the process. The organization is devoted to the collection, propagation, and study of plants in their native habitats, with a potential outcome of selection and eventual introduction. There can be little doubt but that plant diversity--in all its glorious forms--is going to be crucial in keeping the planet habitable, most especially for humans. Peter Del Tredici is a Senior Research Scientist at the Arnold Arboretum. T PAUL MEYER "},{"has_event_date":0,"type":"arnoldia","title":"The Return to China, Mother of Gardens","article_sequence":2,"start_page":4,"end_page":11,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25502","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070af6b.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Meyer, Paul W.","article_content":" The Return to China, Mother of Gardens Paul W. Meyer I n 1929, Ernest Henry Wilson's book China, Mother of Gardens was published, which documented the importance of Chinese plant species to western gardens. Wilson collected plants widely in China between 1899 and 1910. Many of his introductions have become important components of the cultivated flora of our gardens and our cities, and have been used widely in plant hybridization and selection. Wilson's China collections greatly expanded our understanding of the Chinese flora as the richest and most diverse flora in the temperate world and identified the usefulness of many species for cultivation. However, until recently, many of the most important and useful Chinese species in America were the result of limited seed collections, representing only a narrow slice of the genetic diversity and potential of each species. In some cases, all the plants in this country derived from a single plant or a few seedlings. After multiple generations of propagation from seed, symptoms of inbreeding were being observed. Since the 1930s, wars and the political situation in China made it difficult, if not impossible, for western scientists and plant explorers to travel, study, and collect plants in China. However, Chinese botanists were hard at work during this time, cataloguing, describing, and publishing detailed accounts of their flora. With the gradual opening of China following President Nixon's visit in 1972, these publications became more available in western botanical libraries. This new information further documented the richness of the Chinese flora and its potential for further plant exploration, evaluation, and introduction. These data also provided additional information on the natural geographic distribution of species, allowing us to target specific areas for collection in particular parts of their range. By doing so, we could potentially maximize adaptability characteristics such as winter hardiness, heat and drought tolerance, and adaptability to special soil characteristics. Collaboration Breeds Success Earlier expeditions in South Korea, beginning in the 1960s and 1970s, had clearly demonstrated the diversity of plant species still relatively unknown to western horticulture and the importance of studying intraspecific variation and its potential usefulness to landscape crops. Up until this point, collections had been largely one-time efforts and not part of a comprehensive plan to collect over a large geographic range. In the early 1980s, Barry yinger, at that time employed by the United States National Arboretum, proposed a series of Korean collecting trips that would facilitate the collection of specific target taxa over separate geographic and climatic ranges. The resulting trips occurred between 1984 and 1989. They were facilitated by the United States National Arboretum in close collaboration with American and Korean botanical institutions and were highly successful. This collaborative approach based on a multi-year master plan became the model on which we began to build a proposal for a longrange plan for plant exploration in China. Key to the success of the Korean expeditions was the principle of collaboration among institutions as they plan, execute, and follow up on a plant collecting expedition. Most institutions today do not have the financial or human resources to do this work alone. Collaboration allows for the division of responsibilities and of the significant financial commitment needed. It also broadens the range of expertise present in the field. The field work itself is labor Facing page: While on a plant collecting trip for the Arnold Arboretum, E. H. Wilson captured this image of an alpine village (elevation 7,500 feet [2,286 meters]) in the mountains of western Sichuan, China, in August 1908. From the Archives of the Arnold Arboretum. 6 Arnoldia 68\/2 ANTHONy AIEllO reliably cold hardy in USDA Zone 6 (average annual minimum temperature 0 to -10F [-17.8 to -23.3C]). Indeed, seedlings grown have demonstrated superior winter hardiness and, after years of testing, a number of named cultivars have been introduced. Similarly, kousa dogwood (Cornus kousa) was a tried and proven landscape plant but most, if not all, kousa dogwoods in the United States at that time (pre-1980) were descended from a nar row genetic pool. Additional collections made in Korea in the 1980s further demonstrated the great variation within that spe(Left to right) Kris Bachtell, Chris Carley, and Li Jianjun collect ash seeds (Fraxinus cies. By 1990 we were seeing paxiana NACPEC08-016) on a road in Hong He Gu Forest Park, Shaanxi. the promise of greater winter intensive and it is important to have multiple hardiness, increased vigor, and interesting varihands to physically collect the seeds, make and ation in flower bract shape and size from these record the field observations, and complete the collections. As we reviewed Chinese floras we evening tasks of cleaning seed and processing discovered that Chinese botanists had observed herbarium specimens. Most important, coland documented wide variation within this laboration provides multiple sites for propaspecies in China, so much so that they divided gating, growing, and eventually evaluating the what we know as Cornus kousa into multiple collections. Multiple institutions give a variety species. The successes of our Korean collecof testing locales and some level of insurance tions along with the promise of a richness of against seed loss or crop failure. And even if intraspecific variation encouraged us to purall are successful, it allows for a greater genetic sue additional collections of these and other reservoir to be tested and preserved through species in China. permanent living collections. Another specific plant that motivated us was Chinese hemlock (Tsuga chinensis). During the Widening the Pool 1980s and 1990s, hemlock woolly adelgid (AdelSeveral notable collection successes from the ges tsugae) was becoming widespread in the Korean expeditions encouraged us to continue northeastern United States, causing our native this work in China. In 1984, there were two eastern hemlock (Tsuga canadensis) to decline expeditions to islands off the northwest coast of and often die. At both the Morris Arboretum South Korea. The key target species was Cameland the Arnold Arboretum, it was noted that lia japonica. It was believed that populations a Chinese hemlock growing near an infested on these islands would represent the most coldeastern hemlock was resisting infestation. A hardy forms of the species. Seedlings grown few other Chinese hemlocks growing in other in multiple institutions could be selected for arboreta were visited and also were showing hardiness as well as landscape attributes, and resistance. This led to a comprehensive plan ultimately be used in breeding projects, with to re-collect Chinese hemlock from a number the ultimate goal being clones well-adapted and of different locales across its natural range in Return to China 7 China to further study its adelgid resistance and explore horticultural variation and adaptabilities within this species. By the late 1980s, travel to China was becoming more practical and a group of horticulturists who had worked in Korea began to envision a wide-ranging, long-term plan to collaborate with Chinese colleagues. We had already identified a broad geographic arc across northern China that represented areas with a climate parallel to the northeastern United States. We looked at reported mean temperature in both January and July and gave priority to areas with both hot summers and cold winters. Also, we continued to target superior Chinese species already known and grown in the United States that might benefit from new collections and the introduction of greater genetic diversity. We also began the slow process (in the days before email) of establishing contacts with Chinese colleagues and exploring the possibility of meeting with them in China to map a collaborative plan and agreement. Out of this effort, a loose consortium of institutions came together in 1991 to form The North AmericaChina Plant Exploration Consortium (NACPEC). Founding members included lawrence lee of the United NANCy ROSE States National Arboretum (Washington, D.C.), Peter Bristol of the Holden Arboretum (Kirtland, Ohio), and Paul Meyer of the Morris Arboretum of the University of Pennsylvania (Philadelphia, Pennsylvania). NACPEC, the Early Years Our overtures were enthusiastically received and in the autumn of 1991 the founding American members traveled to China to visit a number of botanical and forestry institutions in six cities, and to explore the feasibility of future plant exploration trips. Host Chinese Institutions included the Research Institute of Forestry and Beijing Botanical Garden, the Heilongjiang Academy of Forestry in Harbin, the Chang Chun Forest Botanic Garden, Xian Botanic Garden, and Nanjing Botanic Garden. We visited their herbaria and discussed target species and potential area for collection as well as the nuts and bolts of planning future trips and getting official permissions. With the advice and encouragement of our Chinese colleagues, we laid tentative expedition schedules for the next 5 years. As part of the exchange, NACPEC would assist our Chinese partners by supporting RICHARD SCHUlHOF Kousa dogwood (Cornus kousa) has a broad native range and much variation within the species. Hemlock wooly adelgids (seen as cottony white dots along the branchlets in this photo) attack native eastern hemlock (Tsuga canadensis) but Chinese hemlock (T. chinensis) appears to be resistant. 8 Arnoldia 68\/2 PAUl MEyER PAUl MEyER research projects, facilitating and supporting student and professional exchanges, and by procuring wild-collected and cultivated North American germplasm for evaluation by Chinese institutions. Funding for these expeditions was primarily dependent on the participating NACPEC members' institutions with additional support from the National Plant Germplasm System, a part of the Agriculture Research Service of the United States Department (From right, facing camera) Peter Bristol, Lawrence Lee, and He Lin examine of Agriculture (USDA). The herbarium specimens in the Nanjing Botanical Garden Herbarium on the 1991 USDA recognizes that the pro- planning trip. Research in the herbarium was helpful in pinpointing potential duction of landscape plants sites for future exploration. represents a significant and growing part of American agricultural production and that, in the past, landscape plants were not well represented in the germplasm repository system. Interest in NACPEC grew following the success of these planning efforts. In 1992, longwood Gardens (Kennett Square, Pennsylvania) and the Morton Arboretum (lisle, Illinois), based on their expressed interest and experience, were invited to join NACPEC. later, the Arnold Arboretum (Boston, Massachusetts) and the University of British Columbia In the far north of Heilongjiang, beautiful remnants of the great Manchurian Botanic Garden (Vancouver, forest remain. The tallest trees are Pinus koraiensis, Picea jezoensis, and Abies nephrolepis. Common deciduous species include Betula costata, Betula platyBritish Columbia) joined the phylla var. mandshurica, Fraxinus mandshurica, Acer pictum ssp. mono, and collaboration. Quercus mongolica. The Manchurian forest appears quite similar to northern Planning progressed for the New England forests. first NACPEC full plant exploration trip to the province of Heilongjiang, ters, pole pruners, packing bags, and sphagnum located in the far northeastern corner of China. moss for packing and shipping seed. Once we We were hosted by Professor Jin Tieshan, a arrived, we had to navigate the protocols of renowned professor of forestry at the Heilongjiimportation of supplies and later the exportang Academy for Forestry. This first expedition ing of seed. As with governments everywhere, to China was a great learning experience for these procedures are never fast or easy. the American visitors and our hosts alike. We We were overwhelmed by the commitmailed much of our equipment over in advance, ment and hospitality of our hosts. They did including herbarium presses, papers and bloteverything possible to help us professionally Return to China 9 PAUl MEyER PETER DEl TREDICI and to look after our safety and human comforts. The modern world had not yet arrived in rural Heilongjiang in 1993 and we had a chance to experience the beauty of the traditional agrarian life in northeast China. In the far north of Heilongjiang, we got to see remnants of the once great Manchurian forest with Korean pines (Pinus koraiensis), yezo spruce (Picea jezoensis), and Manchurian fir (Abies nephrolepis) towering well over 100 feet (30.5 meters) tall. By the end of this month- (Left to right) Sheng Ning , a local host, and Jeff Lynch collect seeds of Acer triflolong trip we had collected rum, a maple valued for its exfoliating bark and brilliant red-orange fall color. 112 accessions. Especially notable collections include Maackia amurensis, a potentially useful urban street tree; Pinus koraiensis, a beautiful and fast growing five-needled pine; and Abies holophylla, one of the firs best adapted to areas with hot summers. Where possible, each accession included dried pressed specimens for herbaria of both Chinese and American institutions and seed lots to be grown and evaluated in our institutions. The herbarium specimens serve as an important part of the scientific documentation of each germplasm collection and a perma- This Chinese man collected a bumper harvest of Korean pine (Pinus koraiensis) in the edible seeds (comnent record of the occurrence cones from the local forest in Jilin. After dryingand the sun, Korean pine is the most monly called pine nuts) are extracted, cleaned, packed. of that species in the wild. common source of pine nuts in world commerce because they are relatively large, This work is especially urgent plentiful, and inexpensive. and important today as China is being developed at an unprecedented pace. that first diplomatic trip in 1991, NACPEC Mountainous areas that were largely pristine has sponsored a total of 12 plant collecting in the 1980s were being developed with tourtrips to China. ist resorts and aerial tramways in the 1990s, A World of Opportunities putting increased pressure on the already limThe outcome of this work is hard to fully meaited natural areas in China. The success of sure as it has affected so many individuals and this first expedition energized the team to institutions in so many ways. Over the years, continue planning for two separate expediNACPEC plant explorers have had a chance to tions in 1994 and others in later years. Since Return to China 11 visit and study innumerable Chinese plants in their natural habitat and to learn from Chinese colleagues about the plants' economic and folk uses. By seeing a plant growing in its natural habitat, we can glean insights into the growing conditions to which it is best adapted. It has also given our collectors the opportunity to lecture to groups of professionals and amateurs about the importance of conserving our planet's plant resources. Additionally, NACPEC institutions have hosted many Chinese colleagues and students for study visits and extended internships in the United States over the past 20 years, as part of our broader academic exchanges. Today we are well aware of the dangers of introducing a new invasive species. In many instances we are focused on re-collecting new genetic material of plants that have already proved themselves as well-behaved, handsome landscape plants. When in the field, many potentially invasive plants were left uncollected. Warning signs include an aggressive habit in their natural environment or the existence of related species which have already become unruly in the United States. Back at home, curators keep an eye on plants in the botanic gardens and those showing invasive tendencies are typically removed. With each trip, NACPEC has become increasingly focused on a limited number of target species that address specific emerging needs, such as resistance to the hemlock woolly adelgid or the emerald ash borer, rather than on general collecting. Over the years, hundreds of plants have been shared with NACPEC members and many other non-member institutions. In all, the database of NACPEC collections lists 1,348 accessions with over 6,000 plants in 9 institutions. Each collector may have their favorites, and individual plants are attracting attention as possible cultivar introductions. But beyond the garden merit of these plants, perhaps one day some \"ugly duckling\" may be found to contain genes for resistance to some still unknown virulent disease or insect, or may contain a compound effective in the fight against cancer. No doubt the value of these collections will continue to emerge for decades and maybe even centuries to come in ways we cannot yet imagine. NACPEC is probably the most successful, broad-based, and long-lived collaboration of its sort anywhere in the world. And after nearly 20 years of active collecting in its countryside, modern China continues to indeed be the \"Mother of Gardens.\" Bibliography Aiello, A. S. 2005. Evaluating Cornus kousa cold hardiness. American Nurseryman 201(5): 3239. Aiello, A. S. 2009. Seeking cold-hardy camellias. Arnoldia 67(1): 2030. Aniko, T. 2006. Plant exploration for Longwood Gardens. Timber Press, Portland, Oregon. Del Tredici, P. and A. Kitajima. 2004. Introduction and Cultivation of Chinese Hemlock (Tsuga chinensis) and Its Resistance to Hemlock Wooly Adelgid, Journal of Arboriculture 30(5): 282287. Meyer, P. W. 1999. Plant Collecting Expeditions: A Modern Perspective. The Public Garden 14(2): 37. Meyer, P. W. 2000 Plant Collecting Expeditions: A Modern Perspective. In: J. R. Ault (ed.) Plant Exploration: Protocols for the Present, Concerns for the Future (Symposium Proceedings). Chicago Botanic Garden, Chicago, Illinois. pp. 712. Meyer, P. W. and S. J. Royer. 1998. The North American Plant Collections Consortium. The Public Garden 13(3): 2023. Meyer, P. W. 1985. Botanical riches from afar. Morris Arboretum Newsletter 14(1): 45. yinger, B. 1989. Plant Trek: In pursuit of a hardy camellia. Flower and Garden 33(2): 104106. yinger, B. 1989. Plant Trek: On site with hardy camellias, Sochong Island, Korea. Flower and Garden 33(3): 6266. Paul W. Meyer is the F. Otto Haas Director of the Morris Arboretum of the University of Pennsylvania in Philadelphia, Pennsylvania. Facing page: This gateway building is part of the beautiful Taoist temple complex on the slopes of Wudang Mountain in Hubei. Taoist monks find tranquil spirituality in nature. Over the centuries they have helped protect this botanically rich forest, dubbed \"horticultural heaven\" by NACPEC expedition members. Photo by Paul Meyer. "},{"has_event_date":0,"type":"arnoldia","title":"Sharing and Enjoying the Joint Botanical Expeditions","article_sequence":3,"start_page":12,"end_page":15,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25500","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070ab6d.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Shanan, He","article_content":"Sharing and Enjoying the Joint Botanical Expeditions He Shanan A fter nearly thirty years of isolation from each other, the first modern Sino American botanical expedition was initiated in 1980, with five American botanists participating in a joint expedition to Shennongjia in China's Hubei province. In a reciprocal activity, five Chinese botanists then joined a field expedition in the United States for more than four months in the spring and summer of 1982. It was a historic expedition--the largest Chinese team to make a field trip outside of China in many years--and it generated a sound mutual understanding and cooperative base for the further development of botanizing activities. Some ten years later, a series of joint expeditions began in China, organized by NACPEC and a group of Chinese botanical gardens, which have continued for twenty years already. Improving on the first SinoAmerican botanizing trip, the NACPEC expeditions have covered a very wide range of geographical areas and have had teams composed of members from a number of different disciplines. By all accounts, both the American participants and their Chinese botanical garden hosts have been well satisfied by the efficiency, valuable collections, and the mutually beneficial exchanges of science and technology. Speaking as a member of the Chinese botanical gardens team, I would like to express my great interest and satisfaction in the project, since it has made a considerable contribution to the ex situ conservation collections in Chinese botanical gardens. First, it increased the accessions and enriched the geographical diversity of the botanical gardens' living collections. These collections typically consist of relatively few individuals of a given species, and they are often collected from only a few geographic localities. According to modern concepts of ex situ conservation, a well-balanced germplasm collection should consist of numerous individuals Professor He Shanan in the propagation house at the Nanjing Botanical Garden. from multiple locations. Second, these kinds of collaborative projects can save both money and human resources by sharing plant materials collected with other botanical gardens in different regions, thereby reducing the risk of losing precious plant germplasm. And third, this project provides good opportunities for exchanging scientific information, methodology, and experience. Personally, I am very interested in knowing that most botanical gardens in the United States have the same team doing the collecting activities in the wild, managing the propagation PAUl MEyER Joint Botanical Expeditions 13 HARVARD UNIVERSITy HERBARIA Herbarium specimens like this one of Emmenopterys henryi are permanent scientific records of individual collections that may well last long beyond the living material. In most cases, multiple herbarium specimens were made of each collection for sharing among the Chinese and American institutions. 14 Arnoldia 68\/2 PAUl MEyER Emmenopterys henryi is a rare and endangered tree species native to China. E. H. Wilson first collected this species in 1907 on an expedition for the Arnold Arboretum. He described it as \". . . one of the most strikingly beautiful trees of the Chinese forests, with its flattish to pyramidate corymbs of pure white, rather large flowers and still larger white bracts.\" The 1994 expedition to Wudang Shan found and collected E. henryi in Hubei (top photo).Some of the seeds from this collection went to the Nanjing Botanical Garden, and the resulting seedlings are seen growing in the NBG propagation house (bottom right). An herbarium specimen from this collection (previous page) shows the persistent bracts and oblong seed capsules. In cultivation, Emmenopterys henryi is notorious for taking decades to start blooming, though this precocious specimen (bottom left) at the Quarryhill Botanical Garden in California bloomed at just six years of age. PETER DEl TREDICI JESIKA JENNINGS Joint Botanical Expeditions 15 American and Chinese expedition members shared information and expertise throughout the collection process. in the garden, and taking care of the resulting collections. Such a unified approach encourages botanical garden staff to have strong feelings of attachment to the collections. In Chinese botanical gardens, sometimes these three activities were conducted by different groups of people; for example, the taxonomists and their technicians conducted the expeditions in the wild, and the horticulturists and gardeners propagated the plants and maintained the introduced materials. It is also helpful for Chinese botanical gardens to learn to emphasize and to standardize the record system both in the wild and in the garden. Summarizing the achievements of the cooperative projects, it is obviously very positive, especially as we know that there are so many new plants released. More investigations and more collections are critically important as the planet faces the serious challenge of climate change and ex situ conservation becomes the only effective method for saving plants in the face of relentless urban expansion. I would like to suggest that the NACPEC project should continue its development and move ahead with follow-up research on the plants already collected. Conserve more plants for humanity! Have great success in the future! He Shanan was Director of the Nanjing Botanical Garden in Nanjing, China, from 1983 to 1998. PETER DEl TREDICI "},{"has_event_date":0,"type":"arnoldia","title":"Traveling in China Photo Features","article_sequence":4,"start_page":16,"end_page":19,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25503","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070b36f.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":null,"article_content":"TravelIng In ChIna S Clockwise from upper left: haring narrow rural roads with large logging trucks led to plenty of white-knuckle moments in China (top two photos). A close encounter on the road to Changbai Shan in 1997 turned nearly fatal--by a matter of inches-- when an oncoming truck swerved, directing an unsecured log straight toward the NACPEC expedition van. The end of the log neatly peeled off the driver's door like the lid on a sardine tin (above and at left). Paul Meyer, looking slightly stunned, observes the damage. Ever the plantsmen, the expedition members identified the offending log as Manchurian linden (Tilia mandshurica). Upper left photo Paul Meyer, all others Peter Del Tredici Traveling in China Photo Features 17 O ther travel hazards included muddy roads and various waterways as well as sinkholes and road construction (top two photos). Clockwise from right: This apple vendor used a low-tech but efficient means of transport, a wooden handcart. Once at the collection sites, foot power became the required mode of travel. Charles Tubesing leads the expedition members through a patch of Oplopanax elatus. The rewards of hiking included incredible views of the scenery such as Tianchi (Heavenly) Lake at 2,000 meters (about 6500 feet) elevation in the Changbai Shan. Unfortunately, NACPEC expedition members did not see the mysterious monsters that local legend says inhabit this volcanic crater lake. All photos Paul Meyer C FIeld ColleCTIng ollecting seeds, plants, and herbarium specimens is a team effort on expeditions. Clockwise from upper left: Jeff Lynch and Paul Meyer check out an unknown elm species (Ulmus sp.) on a hill above the Yalu River, which divides China from North Korea (seen in the background). Though woody plants dominate among NACPEC collections, some herbaceous plants such as Paeonia obovata (held by Sheng Ning) have also been collected. This peony grows in moist, fertile soils in the woodland understory. The glamorous work of plant explorers includes picking up Manchurian ash (Fraxinus mandshurica) seeds along a roadside. Out on a limb: Mr. Park, \"the barefoot guide,\" balances precariously while collecting branches from a three-flowered maple (Acer triflorum). Chris Carley, Bai Genlu, and Li Jianjun strip seeds from harvested branches of Acer stachyophyllum ssp. betulifolium. Photos clockwise from upper left: Peter Del Tredici, Paul Meyer, Kris Bachtell, Peter Del Tredici, Anthony Aiello R ecording detailed data is an essential part of the collection process. Clockwise from upper left: In the mountains of the Beijing area, Rick Lewandowski (left) takes notes on location and surrounding flora while Ned Garvey (center) writes out labels for the seeds being packaged by Charles Tubesing (right). Tools of the collecting trade include extendable pole pruners, held here by Wang Xianli. Pole pruners are put to use collecting Fraxinus insularis in the rain during the 2008 expedition. A cluster of fruit collected from Farges filbert (Corylus fargesii). A thin-shelled nut is held within each tightly wrapped involucre. This species was previously rare in the United States and is a valued addition to the USDA's Corylus germplasm collection. Upper right photo Peter Del Tredici, all others Kris Bachtell (Continues on page 36) "},{"has_event_date":0,"type":"arnoldia","title":"By the Numbers: Twenty Years of NACPEC Collections","article_sequence":5,"start_page":20,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25487","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060b36b.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Aiello, Anthony S.; Dosmann, Michael S.","article_content":"By the Numbers: Twenty Years of NACPEC Collections Anthony S. Aiello and Michael S. Dosmann B eginning with the initial feasibility expedition in 1991, NACPEC has conducted a total of 12 botanical expeditions to China (Table 1). These represent a concerted effort to systematically investigate and explore varying climatic areas, habitats, and ecosystems across a wide geographic range. Although quite comparable in land mass, China has much greater plant diversity than the United States. Target areas for NACPEC expeditions were determined based on climate information and cover a broad arc from central China where the Qinling mountain range forms the continental divide separating north and south China (the yellow and yangtze River systems), through the mountain ranges west of Beijing, to the far north and northeast of the country in the provinces bordering Russia and North Korea. Although there is a fascinating and diverse flora in Sichuan and yunnan, these provinces have not been primary targets because plants from this warmer region of China have not performed particularly well in the climates of most NACPEC members. The goals of each NACPEC trip have varied-- on many trips we collected broadly, working from a large list of target plants, while on other trips we focused on specific taxa (e.g., Tsuga in 1998 and 1999, and Fraxinus in 2008). (See map on page 26 for locations covered on each trip.) The contributions from these trips have resulted in a wealth of knowledge about the characteristics and ecology of Chinese plants, represented by copious collection notes and herbarium specimens. And, by bringing germplasm back to North America and integrating it into the living collections spread among all of the members of the consortium, we collectively learn how these individuals respond under cultivation to our diverse growing conditions. The expeditions are summarized in separate trip reports that are housed in the libraries of the participating institutions and generally consist of two parts: a trip journal and the field notes. The journal recounts the daily activities of the trip and also sets out the context for the various plant collections. The detailed field notes provide extensive information on all of the collections for an expedition. These trip reports provide a resource for current and future exploration efforts. Sifting the Statistics As more NACPEC collections were propagated, distributed, and evaluated, it became clear that we should report on the results of the NACPEC expeditions. Our goal with this article is to provide information on the successes and challenges of collecting in this modern era, and to evaluate the significance PETER DEl TREDICI Detailed notes are taken for each collection. Here, Paul Meyer records a collection location from a GPS (global positioning system) device. Sunrise in Xia Ban Si (Cloud Sea), Shaanxi. The photograph was made at an elevation of over 9,000 feet (2,743 meters), the highest point reached on the 2008 expedition. Photo by Anthony Aiello. of NACPEC collections to botany, plant conservation, and ornamental horticulture. A similar case study was published by Dosmann and Del Tredici in their review (2003) of the 1980 SinoAmerican Botanical Expedition (SABE), another collaborative trip that yielded abundant herbarium and germplasm collections from Hubei. We wanted to know if there were any similarities or differences between the SABE and the NACPEC trips, and if there were lessons learned that could be applied not only to future collecting trips, but to living collections management in general. NACPEC members regularly combine and update lists of their expedition holdings into a complete plant survey, and this was most recently completed in late 2009 (for online access to this information see the Data Base of Asian Plants in Cultivation [DAPC] http:\/\/ www.quarryhillbg.org, the BG-Base Multi-Site Search page http:\/\/www.bg-base.com, and the individual institutions' websites). This combined inventory aids curatorial decisions among the collaborating institutions, helps to find missing or unusual collections, and focuses future collecting efforts. We used this combined inventory as the basis for the descriptive statistics provided in this article. These statistics include all of the collections made on the 12 NACPEC expeditions, as well as seeds collected in 1994, 1997, and 2001 by Professor Cui Tiecheng (formerly of the Xi'an Botanic Garden), and a few sets of seeds received as exchanges. living germplasm data came from the NACPEC institutions, plus the holdings at the Dawes Arboretum. A very recent inclusion in this survey is the University of Idaho Arboretum and Botanical Garden, which received a seed distribution following the 1993 Heilongjiang and 1994 Beijing expeditions (see page 24). NACPEC: Who We Are and What We Do S I N C E its inception, NACPEC's efforts have been motivated by a number of goals, including: Broadening the genetic pool of species already in cultivation, with Conserving rare species. Selecting improved ornamental forms. Evaluating and introducing appropriate new species. Increasing our understanding of botanical diversity throughout China. Collaborating with key institutions in the national and international botanical community. particular emphasis on extending cold hardiness and increasing vigor, improving adaptability to stressful environments, and increasing insect and disease resistance. NACPEC consists of eight member institutions plus partner organizations that contribute to the success of our collecting efforts and the wide distribution of valuable germplasm. Each location not only has unique growing conditions that are favorable for certain types of taxa or those from specific parts of China, but the individual missions and collections policies of each institution are novel. Such diversity is truly an asset. The NACPEC members are: The Arnold Arboretum of Harvard University, Boston, MA The Holden Arboretum, Kirtland, OH Longwood Gardens, Kennett Square, PA The Morris Arboretum of the University The Morton Arboretum, Lisle, IL United States National Arboretum, University of British Columbia Botanical USDA Woody Landscape Plant Germplasm Repository, Beltsville, MD Garden, Vancouver, BC Viburnum betulifolium of Pennsylvania, Philadelphia, PA Washington, DC Other partners in these efforts include the Dawes Arboretum in Newark, OH as well as several Chinese botanical institutions listed in Table 1. KRIS BACHTEll Twenty Years of NACPEC Collections 23 What's a Collection? V I S I T O R S to public gardens may associate the word \"collection\" with groups of living plants, often labeled with signs such as \"Maple Collection\" or \"Conifer Collection.\" But on NACPEC expeditions, we use the word collection in a different way. A collection results from one specific act of collecting and may comprise one or more products. Each NACPEC collection receives a unique alpha-numeric code for identification. For example, on the 2005 trip to Gansu we collected seeds from a Cercidiphyllum japonicum and also collected herbarium specimens from the tree. A single collection number, NACPEC05-059, was assigned to both the seeds and herbarium specimens, and any associated data also carry that collection number. Once a NACPEC collection (in the form of seeds, cuttings, plants, or herbarium specimens) arrives at a botanical garden or arboretum, it is typically assigned an accession number unique to that institution. The Cercidiphyllum collected in Gansu is accession AA # 126-2007 at the Arnold Arboretum and MOAR # 2005-192 at the Morris Arboretum, but both institutions can track their accessions back to the original NACPEC collection, which makes ongoing evaluation This Cercidiphyllum japonicum growing at the and reporting easier and more accurate. Any instituArnold Arboretum is identified by the Arnold tion that receives clonally propagated plants of that Arboretum accession number 126-2007-A NACPEC collection in the future may give it their own (letters identify individual specimens within the accession group), but the label information accession number but will also retain the original also includes the original NACPEC collection number, NACPEC05-059. NACPEC collection number in their records. There have been a total of 1,350 unique NACPEC collections since 1991 (Table 2). Of these, 71% (961 collections) are represented by herbarium specimens and 93% (1,250 collections) were originally collected as germplasm (primarily seeds but occasionally seedlings or cuttings); most collections comprised both germplasm and herbarium vouchers. Of the 1,250 germplasm collections made, more than half (56%) are currently represented by living plants among the various member gardens, a percentage somewhat greater than the 1980 SABE (258 of initial 621, or 41%). Certainly, as in the 1980 SABE, an inability to successfully propagate some germplasm collections led to their initial failure. For example, in the case of some taxa such as Acer (maples), seeds collected may be empty and therefore not viable. Similarly, seeds of other taxa may germinate but only grow into weak plants that do not make it out of the propagation\/production phase. And there are also those taxa that make it out onto the grounds only to perform poorly and eventually die. Thus, a \"success rate\" of around 50% is not uncommon. In directly comparing the NACPEC and SABE collections, we wondered what the contributing factors might be for the slightly higher rate among NACPEC collections (56% versus 41%). Certainly, some of the more recent NACPEC collections are still going through the pivotal propagation\/production phase, which contributes to the higher PAUl WARNICK, UNIVERSITy OF IDAHO RICHARD NASKAlI Got NACPEC Plants? W H I L E writing this article, we happened upon information that added significantly to our inventory and the compilation of our statistics. Charles Tubesing, curator at the Holden Arboretum, forwarded to me a newsletter from the University of Idaho Arboretum and Botanical Garden. In that newsletter, Paul Warnick wrote about the development of an arbor to hold vines that they had grown from seeds collected by NACPEC. In further correspondence with Paul, I learned that their institution holds 246 NACPEC plants representing 55 taxa and 64 collections. These include 5 collections A number of NACPEC collection plants grow that previously existed at only one institution, 3 colon the vine arbor at the University of Idaho Arboretum and Botanical Garden, including, lections that previously existed as a single plant at a Clematis mandshurica HLJ-073 and Vitis lone institution, and 2 collections that we previously amurensis BJG-039, top, and Actinidia arguta BJG-025 (male flowers), bottom. thought were dead altogether. While we knew that NACPEC collections had been distributed far and wide, this one instance illustrated just how pivotal distributions outside the NACPEC network can be. In light of this information, we would be very interested in hearing if any other organizations have NACPEC collections in their gardens. If so, please contact Anthony Aiello at aiello@upenn.edu . We would be happy to include your records in future NACPEC inventories. --ASA success rate. However, we believe that the overwhelming reason for the greater success is due to the unique nature of NACPEC: wide initial distribution of germplasm--and continuing distribution of surplus plants and vegetative propagules--to a network of gardens and arboreta with unique growing environments. lighty (2000) described several barriers to successful expeditions, one of them being the \"too-rapid rate of entry of plants into the system\" that then overwhelms staff and facilities. Because of its distributive and collaborative nature, NACPEC may have found a way to break this barrier. Notable Successes There are some remarkable success stories from the trips of the early 1990s, with 62% of the 1993 Heilongjiang collections and 69% of the 1994 Beijing expedition plants still alive today. What might have contributed to these successes? Germplasm from these trips was widely distributed to numerous NACPEC institutions, and by and large it was well-adapted to these varying climates. And even though the 1999 trip to Sichuan has a fairly low percentage of living germplasm (40%), significant collections resulted from this expedition, most notably Tsuga chinensis var. oblingisquamata (Table 3). Another important statistic revealed in our assessment is the level of duplication among collections. Of those unique germplasm collections that are currently alive, approximately 60% grow in at least two different institutions. This duplication insures against the loss of valuable material and also provides opportunities for broader evaluation and study across a number of different growing sites. The numbers do not always tell the whole story, however. For example, there are only 15 living collections (out of an initial 30) from the 1995 Collaborative distribution of seed and plant collections helps prevent individual NACPEC member institutions from reaching \"propagation overload.\" ERIC lA FOUNTAINE Chimonanthus praecox (the cultivar `Grandiflorus' is seen here) bears fragrant yellow flowers in late winter or early spring. NANCy ROSE 26 Arnoldia 68\/2 1 Heilongjiang 1 Inner Mongolia A.R. Liaoning Jilin 6 BEIJING Ningxia 2 Tianjin Shanxi Hebei Shandong Qinghai Gansu 9 5 Shaanxi 10 4 5 11 3 Henan Anhui Hubei Jiangsu 8 Sichuan Chongqing 7 Jiangxi SHANGHAI Zhejiang Hunan Guizhou 7 7 Fujian Guangdong Macau S.A.R. Hong Kong S.A.R. Yunnan Guangxi Zhuang A.R. 0 250 125 250 500 1,000 Kilometers 500 Miles Hainan 0 NACPEC EXPEDITION LOCATIONS 1 1993Heilongjiang 2 1994Beijing 3 1994Hubei 4 1995Shaanxi 5 1996ShaanxiandGansu (QinlingMountains) 6 1997Jilin(ChangbaiShan) 7 1998Anhui,Guangxi,andJiangxi 8 1999Sichuan 9 2002Shanxi 10 2005Gansu 11 2008Shaanxi Twenty Years of NACPEC Collections 27 Shaanxi expedition. One of these is Chimonanthus praecox (SHX033), a shrub noteworthy for its fragrant flowers in early spring. This plant is well known horticulturally, but as far as we can determine, this is the only wild-collected collection in North America. It is represented by only five plants at the Morris Arboretum and is an example of a collection growing at only one institution. This type of collection, held at only one institution, illustrates one of the challenges facing NACPEC as well as others engaged in germplasm acquisition. Forty percent of the NACPEC collections grow in just a single place. Even though most of these exist as multiple plants (as in the Chimonanthus), they are potentially at risk and need to be prioritized for distribution. But at even greater risk are those collections that exist as just a single plant in a single institution. For the NACPEC collections, 15% fall into this category. While they are clearly the most tenuous collections and the highest targets for propagation and distribution, this is considerably lower than the 45% of single-plant collections surviving from the SABE. Again, the unique collaborative and distributive nature of NACPEC contributes to this lower number. The annual inventory of combined holdings is the first step in alerting NACPEC members of the rarity of their own holdings. A P R I M E example in the category of a single plant is Magnolia biondii (QLG062A) collected in 1996 in the Foping Nature preserve in Shaanxi. This rare magnolia is a close relative of Magnolia stellata and is growing as a single individual at the Morris Arboretum. In light of its high conservation value and its rarity in cultivation, it becomes a very important target for propagation and distribution to other botanic gardens. PHIlIPPE DE SPOElBERCH KOEN CAMElBEKE These photographs show the distinctive long carpel (in center of flower) and long fruit structure with red-ariled seeds of Magnolia biondii specimens growing at the Arboretum Wespelaar in Belgium. 28 Arnoldia 68\/2 PAUl MEyER In contrast to these sparsely represented taxa, there are a number of collections that are widely held among the NACPEC members. There are 13 collections grown in at least seven institutions, making them ideal candidates for further evaluation for broad adaptability as well as uniformity. Topping this list is Acer pictum ssp. mono BJG141, held by nine institutions, followed by Acer davidii ssp. grosseri (BJG017) and Corylus fargesii (QlG231), each held at eight gardens. It is difficult to know why some plants are grown more successfully and widely compared to others, but factors include initial quantities, broad original distribution, seed viability, curatorial interest, and broad adaptability to an array of growing conditions. In combined NACPEC holdings, Acer is the most frequently collected genus--not surprising given that China is the center of diversity for maples, with 99 of the 129 species worldwide occurring there. Keen member interest in Acer (five of the NACPEC members and partners--the Arnold, Dawes, Morris, and Morton Arboreta, and UBC Botanic Garden--are members of the North American Plant Collections Consortium's multi-site Acer collection), combined with the great natural diversity has led to the A large specimen of Acer pictum ssp. mono growing in Heilongjiang. extensive holdings in this genus. There have been 106 distinct collections of griseum, A. miyabei ssp. miaotaiense, and maple, representing 33 taxa. Of these, 73 (10% A. yui. These collections represent a signifiof all living NACPEC collections) are represencant increase in the diversity of maples collted by living germplasm for a total 29 taxa and ectively held not only by NACPEC members, 585 plants among all of the member institubut among other North American institutions tions. Among these are garden-worthy plants due to redistribution. such as Acer davidii (including ssp. grosseri), After Acer, other frequently collected taxa A. pictum ssp. mono, and A. triflorum; plants include Viburnum (20 taxa), Quercus and rarely grown in North American botanic Euonymus (15 each), and Rhododendron (13). gardens such A. ceriferum and A. sterculaceum Of special significance are the 26 total collecssp. franchetti (A. tsinglingense); and plants tions representing 11 taxa of Fraxinus. Some of high conservation value, including A. of these ash taxa, such as F. insularis and F. PAUl MEyER Upper left, Rick Lewandowski measures the diameter of an impressively large specimen of paperbark maple (Acer griseum); upper right, Acer ceriferum growing in the wild in China; lower, A cultivated specimen of three-flowered maple (Acer triflorum). paxiana, may eventually hold the key to solving the emerald ash borer epidemic that is devastating native and planted populations of North American ash species. Tsuga is another genus that stands out among the collections, not because of the diversity of taxa but for the number of collections made. Representing a classic case of a genetic bottleneck, T. chinensis had been introduced as a single individual into the United States in 1910. Starting in the 1990s, Tsuga chinensis became a prime target for NACPEC collecting expeditions because of the immediate threat to North American hemlocks by hemlock wooly adelgid (Adelges tsugae). A total of 33 collec- NANCy ROSE ANTHONy AIEllO Chinese hemlock (Tsuga chinensis) in its native habitat. Photo by Kris Bachtell. tions of the adelgid-resistant T. chinensis and its varieties were made from across its native range, 19 of which are represented by living plants (Table 3). The original collections-- mostly by seed but in some cases as seedlings or even cuttings--were made from several provinces from southeast China through the northwestern limit of its range in southern Gansu. Of the 18 seed collections, 17 are represented by at least one plant; none of the cuttings and few of the seedling collections are extant. Because of their high priority, these plants have been widely distributed among the NACPEC members and show that targeted collecting (instead of broad, opportunistic collecting) can greatly increase the diversity of germplasm among our collective holdings. These plants have been widely distributed among other North American botanic gardens, aiding in research on and wider introduction of Chinese hemlock. What Have We Learned? After nearly 20 years of collecting germplasm and herbarium specimens, we can draw a number of important conclusions. Overall these statistics point out the importance of collaboration in sustaining the NACPEC collections. Without the combined efforts of the member organizations, it is difficult to imagine how these expeditions would have occurred, let alone how the plants would have been subsequently propagated and maintained over a period of time. In total the herbarium specimens and living collections represent material of horticultural, botanical, and conservation significance. The sum of the consortium work is certainly greater than its individual parts, leading to significant scientific contribution and a deeper understanding of the Chinese flora as well as its horticultural potential. This is an important point to emphasize--NACPEC's goals are broader than simply introducing gardenworthy plants. First and foremost is the primary scientific documentation of botanical diversity. Additional lessons learned include the need for sustained and repeated collecting within varied geographic, climatic, and ecological ranges, and the importance of vision and long-term planning. looking forward, this analysis will prove a useful tool as we focus on additional Table 1. List of NACPEC expeditions, abbreviations, participants and dates. Trip Name 1991 Initial Feasibility Expedition (Beijing, Heilongjiang, Jilin, Shaanxi, Jiangsu) 1993 Expedition to Heilongjiang Abbreviation LL Participants Bristol, Peter; Holden Arboretum Lee, Lawrence; U.S. National Arboretum Meyer, Paul; Morris Arboretum Bachtell, Kris; Morton Arboretum Bristol, Peter; Holden Arboretum Meyer, Paul; Morris Arboretum Gao Shi Xin; Heilongjiang Academy of Forestry Jin Tae Shan; Heilongjiang Academy of Forestry Liu Jun; Heilongjiang Academy of Forestry Bachtell, Kris; ; Morton Arboretum Lewandowski, Rick; Morris Arboretum Garvey, Edward; U.S. National Arboretum Tubesing, Charles; Holden Arboretum Liu Mingwang; Beijing Botanical Garden IBCAS Conrad, Kevin; U.S. National Arboretum Del Tredici, Peter; Arnold Arboretum Meyer, Paul W.; Morris Arboretum Thomas, R. William; Longwood Gardens Hao Riming; Nanjing Botanic Garden Mao Cailaing; Nanjing Botanic Garden Garvey, Edward; U.S. National Arboretum Lewandowski, Rick; Morris Arboretum Cui Tiecheng; Xi'an Botanic Garden Ault, James; Longwood Gardens Conrad, Kevin; U.S. National Arboretum Lewandowski, Rick; Morris Arboretum Kim Kunso; Norfolk Botanical Gardens Cui Tiecheng; Xi'an Botanic Garden Bachtell, Kris; Morton Arboretum Del Tredici, Peter; Arnold Arboretum Lynch, Jeffrey; Longwood Gardens Meyer, Paul W.; Morris Arboretum Tubesing, Charles; Holden Arboretum Wang Xian Li; Shenyang Institute of Applied Ecology Cao Wei; Shenyang Institute of Applied Ecology Sheng Ning; Nanjing Botanical Garden Lewandowski, Rick; Morris Arboretum Garvey, Edward; U.S. National Arboretum Li Weilin; Nanjing Botanical Garden Wang Qing; Nanjing Botanical Garden Belt, Shawn; U.S. National Arboretum Garvey, Edward; U.S. National Arboretum Stites, Jerry; Longwood Gardens Wang Qing; Nanjing Botanical Garden Aiello, Anthony; Morris Arboretum Bachtell, Kris; Morton Arboretum Bordelon, Carole; U.S. National Arboretum Bristol, Peter; Holden Arboretum (Chicago Botanic Garden) Tang Yudan; Beijing Botanical Garden IBCAS Aiello, Anthony; Morris Arboretum Bachtell, Kris; Morton Arboretum Scanlon, Martin; U.S. National Arboretum Wang Kang; Beijing Botanical Garden Sun Xue-gang; Forestry College of Gansu Agricultural University Aiello, Anthony; Morris Arboretum Bachtell, Kris; Morton Arboretum Carley, Chris; U.S. National Arboretum Wang Kang; Beijing Botanical Garden Dates 10 Oct 2 Nov 1991 HLJ 25 Aug 28 Sep 1993 1994 Expedition to Beijing BJG 13 Sep 3 Oct 1994 1994 Expedition to Hubei WD 6 Sep 11 Oct 1994 1995 Expedition to Shaanxi SHX 31 Mar 17 Apr 1995 1996 Expedition to Shaanxi & Gansu (Qinling Mountains) QLG 30 Aug 18 Oct 1996 1997 Expedition to Changbai Shan (Jilin) NACPEC97 (CBS) 25 Aug 27 Sep 1997 1998 Expedition to E. & SE. China (Anhui, Guangxi, Jiangxi) 1999 Expedition to Sichuan NACPEC98 (TS98) 5 Oct 22 Oct 1998 NACPEC99 (TS99) 1 Oct 20 1999 2002 Expedition to Shanxi NACPEC02 9 30 Sep 2002 2005 Expedition to Gansu NACPEC05 14 Sep 12 Oct 2005 2008 Expedition to Shaanxi NACPEC08 16 Sep 8 Oct 2008 NANCy ROSE Acer davidii is much admired for its striped bark. This specimen at the Arnold Arboretum (AA#666-94-A) was grown from seed collected during the 1994 expedition to Hubei (NACPEC collection #WD 040). Table 2. Collections totals for formal NACPEC expeditions and other associated collections. Trip abbreviations follow Table 1. TRIP Year Total Herbarium collections # % 0 96 130 149 0 235 122 0 29 71 85 44 961 0 961 0 86 90 77 0 89 85 0 88 91 94 86 79 0 71 Germplasm # 24 102 138 171 30 234 139 42 27 71 85 45 1108 142 1250 % 96 91 96 88 91 89 97 100 82 91 94 88 91 100 93 Living germplasm # % 9 63 95 83 15 120 91 19 11 48 60 43 657 47 706 38 62 69 49 50 51 65 45 41 68 71 96 59 33 56 Exists at only one institution* # % 5 20 26 40 11 48 22 17 6 20 29 20 264 36 294 56 32 27 48 73 40 24 89 55 42 48 47 40 77 42 Exists as a single plant** # % 3 5 11 21 5 22 7 11 1 2 4 4 96 15 109 33 8 12 25 33 18 8 58 9 4 7 9 15 32 15 LL HLG BJG WD SHX QLG NACPEC 97 TS 98 TS 99 NACPEC 2002 NACPEC 2005 NACPEC 2008 Expedition total Other*** Grand Total 1991 1993 1994 1994 1995 1996 1997 1998 1999 2002 2005 2008 25 112 144 194 33 263 143 42 33 78 90 51 1208 142 1350 * A germplasm collection that exists at only one institution; may be of one or multiple plants ** A germplasm collection that is represented by a lone plant at one institution *** Include collections made during tourist visits, as well as those collections made through contract NANCy ROSE Acer pseudosieboldianum is another garden-worthy small maple collected on several of the NACPEC expeditions. 34 Arnoldia 68\/2 CHRIS CARlEy Bai Genlu (back to camera), Wang Kang (white hat), Li Jianjun, and Anthony Aiello examine Acer caesium ssp. giraldii (NACPEC08-014) in Hong He Gu Forest Park, Shaanxi. collecting efforts, and will allow us to focus our efforts on propagating and distributing plants of horticultural, botanical, and conservation significance. lastly, and perhaps most importantly, we acknowledge the cultural exchange that has occurred among the American and Chinese institutions and the individuals involved. Without these lasting relationships none of these expeditions or the resultant collections would have occurred. The end result is a mutual affinity and deep appreciation for the relationships that have ensued. Bibliography Aiello, A.S. 2006. Plant collecting on the eaves of the world. The Plantsman 5 (4): 220225. Aniko, T. 2006. Plant exploration for Longwood Gardens. Timber Press, Portland, Oregon. Del Tredici, P., P. Meyer, R. Hao, C. Mao, K. Conrad, and R. W. Thomas. Plant Collecting on Wudang Shan. Arnoldia 55(1): 1220. Dosmann, M. and P. Del Tredici. 2003. Plant Introduction, Distribution, and Survival: A Case Study of the 1980 SinoAmerican Botanical Expedition. BioScience 53 (6): 588597. lighty, R.W. 2000. An assessment of ornamental plant introduction in the not-for-profit sector. In: J. R. Ault (ed.) Plant Exploration: Protocols for the Present, Concerns for the Future (Symposium Proceedings). Chicago Botanic Garden, Chicago, Illinois. pp. 1422. Meyer, P. W. 1999. Plant Collecting Expeditions: A Modern Perspective. The Public Garden 14 (2): 37. Anthony S. Aiello is the Gayle E. Maloney Director of Horticulture and Curator at the Morris Arboretum of the University of Pennsylvania, and Michael Dosmann is Curator of living Collections at the Arnold Arboretum. Table 3. List of NACPEC Tsuga collections. Taxon Coll. # How Material was Collected Province of origin Collection alive? Number of Institutions growing this collection 4 4 3 4 3 6 7 6 2 3 1 1 Total plants among all institutions 24 26 12 12 7 20 18 23 9 17 1 5 174 4 5 2 3 4 0 22 11 22 9 9 0 73 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 0 0 0 0 0 0 0 0 0 0 3 250 Tsuga chinensis Cui 97-053 Cui 97-054 NACPEC05022 NACPEC05063 QLG013 QLG188 QLG190 QLG193 QLG216 QLG217 SHX017 XBG s.n. Seed Seed Seed Seed Seed Seed Seed Seed Seed Seed Seedling Seed Shaanxi Shaanxi Gansu Gansu Shaanxi Shaanxi Shaanxi Shaanxi Shaanxi Shaanxi Shaanxi Shaanxi yes yes yes yes yes yes yes yes yes yes yes yes 12 living (100%) Subtotal Tsuga chinensis var. oblongisquamata 12 TS 99-018 TS 99-022 TS 99-025 TS 99-026 TS 99-027 TS 99-033 Seed Seed Seed Seed Seed Seed Sichuan Sichuan Sichuan Sichuan Sichuan Sichuan yes yes yes yes yes no 5 living (83.3%) Subtotal Tsuga chinensis var. tchekiangensis 6 TS 98-046F TS 98-058B TS 98-035E TS 98-036E TS 98-040H TS 98-042C TS 98-043E TS 98-044F TS 98-051B TS 98-051C TS 98-052B TS 98-062B TS 98-066B TS 98-066C TS 98-069 Seedling Seed Seedling Seedling Seedling Seedling Seedling Seedling Seedling Cuttings Seedling Seedling Seedling Cuttings Seedling Jiangxi Guangxi Jiangxi Jiangxi Jiangxi Jiangxi Jiangxi Jiangxi Guangxi Guangxi Guangxi Guangxi Guangxi Guangxi Zhejiang yes yes no no no no no no no no no no no no no 2 living (13.3%) 19 living (57.6%) Subtotal Grand Total 15 33 "},{"has_event_date":0,"type":"arnoldia","title":"Traveling in China Photo Features 2","article_sequence":6,"start_page":36,"end_page":39,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25504","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070b728.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":null,"article_content":"36 Arnoldia 68\/2 (Continued from page 19) F ProCessIng reshly collected seeds and herbarium specimens must be processed promptly to prevent spoilage. These tasks occupied many evenings for all of the expedition members. Clockwise from upper left: Soft-fleshed fruits are soaked and sieved to remove any pulp before the seeds are dried and packed. Rick Lewandowski cleans seeds at a trough sink during the 1994 expedition to the Beijing region. Pressed herbarium specimens must be dried quickly or else mold may set in. During a rainy spell, Peter Del Tredici and Mao Cailaing resorted to burning charcoal under the herbarium press to help dry the specimens. Air-drying seeds and herbarium samples. Kris Bachtell and Martin Scanlon begin processing cones of Tsuga chinensis. Dry fruits like these maple samaras are winnowed, carefully cleaned, and sorted before they are packed in labeled bags. Photos clockwise from upper left: Kris Bachtell, Paul Meyer, Peter Del Tredici, Anthony Aiello, Kris Bachtell Traveling in China Photo Features 37 E Food xpedition members enjoyed many elements of Chinese cuisine on their travels. Clockwise from upper left: A vendor at a market in Harbin sells fruit including round Asian pears native to this region and warty, orange, ripened bitter melons. Graduate student Zhang Wei and an expedition driver, Mr. Liu, eat spicy noodles in a restaurant in Min Xian, Gansu. This noodle vendor in Xi'an served up bowls of piping-hot seasoned noodles. In Shanxi's Pangquangou National Preserve, this \"mushroom lady\" was collecting a type of edible mushroom known to grow on Populus davidiana. Photos clockwise from upper left: Paul Meyer, Kris Bachtell, Anthony Aiello, Kris Bachtell B geTTIng To Know ChIna eyond collecting plants, the NACPEC expeditions have provided an opportunity for participants to see and learn more about China's people and landscapes. Clockwise from top: Tall, blond expedition members Bill Thomas and Paul Meyer attracted a lot of attention in the streets of Wudang Shan City, Hubei. This Tibetan woman was harvesting Anemone tomentosa plants to feed to her pigs. Chinese colleagues hosted a birthday celebration for Kris Bachtell in September 2005 (left to right: Anthony Aiello, Kris Bachtell, Zhang Zuoshuang, He Shanan, and Zhang Aoluo and his wife). Opposite page, clockwise from upper left: Peter Del Tredici shoots pool with the locals on a street in the Wudang Shan area, Hubei. Children give pig riding a try, to the amusement of onlookers in a small village in the Mo Gou Forest area in Gansu. A misty view of the Seven Sisters, a set of peaks in the Tai Bai Shan reserve in Shaanxi. Photos this page, clockwise from top: Peter Del Tredici, Kris Bachtell, Kang Wang Photos opposite page, clockwise from upper left: Paul Meyer, Kris Bachtell, Anthony Aiello Traveling in China Photo Features 39 "},{"has_event_date":0,"type":"arnoldia","title":"Planning Future NACPEC Plant Exploration: Challenges and Opportunities","article_sequence":7,"start_page":40,"end_page":47,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25498","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070a76f.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Kim, Kunso; Bachtell, Kris R.; Wang, Kang","article_content":"Planning Future NACPEC Plant Exploration: Challenges and Opportunities Kunso Kim, Kris Bachtell, and Kang Wang T he North AmericaChina Plant Exploration Consortium (NACPEC) is an innovative partnership formed in 1991 between American and Chinese institutions to organize and conduct plant explorations in China. Since its inception, this collaborative effort has been successful in collecting many botanically and horticulturally important plants including paperbark maple (Acer griseum), Farges filbert (Corylus fargesii), and Chinese stewartia (Stewartia sinensis). During the two decades of its existence, NACPEC has witnessed dramatic changes occurring around the globe that have profoundly affected the consortium's collecting efforts, most notably the Convention on Biological Diversity, the complexity of ex situ plant conservation, the impacts of climate change, the spread of invasive species, and the tightening of import\/export regulations. These issues have challenged us to rethink the way we plan and organize plant explorations as the consortium moves forward into the next decade. CURRENT ISSUES AND CHALLENGES Convention on Biological Diversity The Convention on Biological Diversity (CBD) is an international treaty signed at the United Nations Conference on Environment and Development (also called the \"Earth Summit\") in Rio de Janeiro, Brazil, in 1992. The key component relevant to NACPEC activities is Article 15, Access to Genetic Resources (CBD 1999). This provision endorses the sovereign rights of countries over their biological resources. The article not only offers countries an opportunity to revamp their efforts in conservation of biodiversity and sustainable uses, but also gives them leverage over their natural resources. In essence, the possibility of con- ducting any plant exploration in China rests upon consent from the host country based on mutually agreed terms. As a consequence, the consortium may see increasing restrictions imposed by the Chinese government on certain collecting areas including national parks and reserves that are often biologically rich. Described in the same article is another provision called \"Access and Benefit Sharing\" that requires the consortium to establish a clear understanding of how the benefits should be shared. Participating countries are required to share with the host country any benefits arising from commercialization or other utilization of the genetic resources. In the past, NACPEC has made good faith efforts to honor this provision by training Chinese students in plant curation and database management, hosting a number of Chinese scientists during their extended visits to the United States, supporting BG-Base and BG-Map at the Beijing Botanical Garden (IBCAS), supporting field work inside China by a number of Chinese botanists, and by sharing a portion of the royalties from the sale of plants through the Chicagoland Grows Plant Introduction Program. Ex Situ Plant Conservation Conserving rare species has always been one of the objectives in the consortium's collecting trips (Meyer 2000). Despite the high priority given to horticulturally important plants, the consortium's annual inventory survey includes 52 accessions that are on the current IUCN Red List of Threatened Species. Should future explorations emphasize collecting red-listed plants as the major goal? As habitat loss in China continues to increase and more public gardens are responding to the plea to participate in ex situ Facing page top: Farges filbert (Corylus fargesii), a promising tree species for landscape use, was collected in this river terrace habitat in Gansu. Bottom: A temple within the Lingkongshan Chinese pine (Pinus tabuliformis) national preserve, one of a number of botanically rich national preserves in China. Photos by Kris Bachtell. 42 Arnoldia 68\/2 When human development encroaches on the habitat of threatened plant species, ex situ conservation may be the key to preventing species extinction. Here, the \"suburbs\" of a village in Hubei expand up a steep terraced slope. Photo by Peter Del Tredici. plant conservation, there is an opportunity for the consortium to renew its efforts and play a more active role in acquiring threatened species and sampling species from different populations to get a better representation of genetic diversity. The consortium is uniquely positioned to expand its ex situ conservation role. It has established successful collaborations with Chinese institutions, gained invaluable field experience, and established a high standard of documentation that holds important information for breeding, evaluation, and conservation purposes (Bachtell 2000). Capitalizing on its experience and collaborations, future explorations can contribute to advancing the goal of preserving 60% of threatened plant species in accessible ex situ collections as laid out in the Global Strategy for Plant Conservation Target 8 (CBD 2005). Future plant exploration with conservation in mind will need to priori- Some widely grown landscape plants of Asian origin have become invasive in parts of North America. One example is winged euonymus (Euonymus alatus), a popular shrub also known as burning bush because of its bright carmine red fall color. ROBERT MAyER Future NACPEC Plant Exploration 43 tize species by conservation concern and sample multiple populations in order to maximize genetic diversity. When sampling from within populations, care must be taken to collect sufficient germplasm to maximize the capture of genetic diversity for long-term seed storage, or clonal stands for taxa that have recalcitrant seeds. Introducing Horticulturally Superior Plants This was at the heart of initial NACPEC plant exploration efforts and remains an important goal for the consortium. The definition of a superior plant in the early days of NACPEC plant exploration often meant that a plant should have increased cold hardiness, tolerance to stressful conditions (including urban landscapes), and resistance to pests and diseases. Exciting plants with such traits are in the process of being introduced and others are being developed through breeding and selection programs utilizing the rich germplasm of plants the consortium has assembled. While the above criteria are still valid, biological invasiveness has emerged as an area of concern. The need to screen introduced plants for non-invasive characteristics has become a high priority for the consortium, which has stepped up its efforts by excluding from expedition target taxa lists any potentially invasive species. At the same time, participating gardens need to develop vigorous risk assessment protocols to determine the invasive potential of lesser known species from the pool of plants that are maintained in living collections before they reach reproductive stage. Use of reliable predictive modeling to assess invasive risk can allow the consortium to avoid time-consuming screening and expensive processing in the field (Widrlechner 2009). Climate Change Climate change has a huge implication for biodiversity and consequently on the consortium's future collecting initiatives. Major vegetation PAUl MEyER Large expanses of forest in China have been cut down and burned to make way for ginseng cultivation. 44 Arnoldia 68\/2 PETER DEl TREDICI PETER DEl TREDICI Ginseng requires shade to grow, so the forests are replaced with vast expanses of low shade structures, seen here in the Changbai Shan area. shifts are predicted by various climatic models. Hawkins (2008) states three different possibilities: (1) some plants will adapt to new climate conditions through selection or plasticity; (2) some plants will move to higher latitudes or altitudes; or (3) other plants may become extinct. It is predicted that continued climate change will ultimately drive many plants to extinction. An average world temperature rise of 2 to 3C over the next 100 years will result in up to 50% of the 400,000 or so higher plants being threatened with extinction (Hawkins 2008). How should NACPEC's future plant explorations respond to climate change? Studying climate analogues has provided the most useful information to NACPEC in planning the potential target areas in China. The consortium considers seasonal rainfall, mean seasonal temperatures, and summer high and winter low temperatures to identify the target areas (Meyer 2000). Will it be necessary for NACPEC to reconsider expanding the collecting zones towards warmer regions or lower altitudes and latitudes in response to anticipated vegetation shifts? Import\/Export Regulations NACPEC members who participated in past plant explorations are familiar with the complexity of obtaining collecting permits in China. Compliance with rules governing col- Emerald ash borer (Agrilus planipennis). Photo by David Cappaert, Michigan State University, Bugwood.org. Future NACPEC Plant Exploration 45 The 2008 NACPEC expedition focused on collecting Chinese ash species. This large specimen of Fraxinus insularis grows near a country house in Shaanxi. Photo by Kris Bachtell. lecting permits and germplasm importation is absolutely necessary, yet it is one of the most cumbersome aspects of any plant exploration. With the CBD recognition of countries' sovereign rights over their natural resources, it is possible that the consortium will see more limited issuance of collecting permits and increased restrictions on collecting in special areas. Adding to the challenges, there are new rules on importation of plant germplasm into the United States. Even clean seeds with phytosanitary certificates attached do not guarantee an easy entry. Some people mistakenly think that the possession of an import permit issued by USDA APHIS for small lots of seeds means free passage, but it only eliminates the requirement to obtain a phytosanitary certificate from the exporting country (USDA 2008). In view of recent insect and disease outbreaks such as emerald ash borer (Agrilus planipennis), Asian longhorned beetle (Anoplophora glabripennis), and sudden oak death (Phytophthora ramorum), it is understandable that the USDA is increasingly tightening the rules. The threat posed by invasive plant species is another issue that has contributed to stricter importation rules. STRATEGIC PLANNING FOR FUTURE COLLECTING EFFORTS Past plant exploration efforts focused on collecting a broad range of species and a large number of accessions. Priority taxa lists are carefully developed during the trip planning process, but inevitably non-target taxa are added to the list 46 Arnoldia 68\/2 along the way, resulting in increased collections. This practice of collecting a broad range of species will need to change. The current issues and challenges described above necessitate that future plant explorations become more sharply focused. This can be accomplished by the annual survey inventory which examines total living accessions accumulated over the last two decades and shows which institutions hold what accessions. The survey results have helped NACPEC understand gaps in its collections and will help with planning future trips. Based on an analysis of past results, a trip could target a single genus or a few genera depending on the purpose or priority of the taxa. More focused collecting trips allow the initiation of new research projects in response to rapidly changing environmental conditions such as those posed by invasive species. The 2008 Shaanxi expedition was an example of a goal-driven and therefore more focused collecting trip. This USDA-funded trip was specifically for collecting ash (Fraxinus) species that are not well represented in American botanic gardens and arboreta. A percentage of the seeds collected were seed banked within the USDA's National Plant Germplasm System. The resulting progenies are to be incorporated into a feeding preference study to test the ash species' resistance to the emerald ash borer. sortium to focus future plant explorations on collecting a more narrowly defined list of target plants, paying particular attention to preventing accidental introduction of plants with invasive potential. NACPEC has succeeded largely through the close collaborative work between the American and Chinese member institutions. The issues listed above present opportunities for NACPEC to help meet the challenges presented by our changing world. Literature cited: Bachtell, K. 2000. Documenting your collections. In: J. R. Ault (ed.) Plant Exploration: Protocols for the Present, Concerns for the Future (Symposium Proceedings). Chicago Botanic Garden, Chicago, Illinois. pp. 5257. Convention on Biological Diversity. (2 November, 1999). Article 15. Access to Genetic Resources http:\/\/www.cbd.int\/convention\/articles. shtml?a=cbd-15 Convention on Biological Diversity. (27 July, 2005). Global Strategy. Target 8. http:\/\/www.cbd.int\/ gspc\/future.shtml#8 Hawkins, B., S. Sharrock, and K. Havens. 2008. Plants and climate change: which future? BGCI, Surrey, UK. Meyer, P. W. 2000 Plant Collecting Expeditions: A Modern Perspective. In: J. R. Ault (ed.) Plant Exploration: Protocols for the Present, Concerns for the Future (Symposium Proceedings). Chicago Botanic Garden, Chicago, Illinois. pp. 712. USDA. APHIS: Plant Import. (14 July, 2008). On small lots of seeds. http:\/\/www.aphis.usda.gov\/import_ export\/plants\/plant_imports\/smalllots_seed. shtml Widrlechner, M. P., J. R. Thompson, E. J. Kapler, K. Kordecki, P. M. Dixon, G. Gates. 2009. A Test of Four Models to Predict the Risk of Naturalization of Non-native Woody Plants in the Chicago Region. Journal of Environmental Horticulture. 27(4): 241250. Kunso Kim is Head of Collections and Curator, and Kris Bachtell is Vice President of Collections and Facilities, both at the Morton Arboretum in lisle, Illinois. Kang Wang is a Research Horticulturist at the Beijing Botanical Garden in Beijing, China. TOWARDS GREATER COLLABORATION Modern day plant explorers are facing a different set of challenges than did earlier plant explorers. The impact of climate change on vegetation is one obvious reason for the consortium to take a more active role in conserving plants. The CBD has helped focus our attention on how to balance the needs for access to genetic resources and benefit sharing, with the ultimate goal of conservation of biological resources through preservation and sustainable use. In spite of the tremendous success NACPEC has experienced in collecting and introducing plants, there is a need for the con- Facing page: A view across the Wudang Mountain range from atop Wudang Mountain in Hubei. The building in the foreground is part of the Taoist monastery and temple complex for which this mountain is famous. Photo by Paul Meyer. "},{"has_event_date":0,"type":"arnoldia","title":"Paperbark Maple Acer Griseum","article_sequence":8,"start_page":48,"end_page":50,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25497","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070a36b.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Meyer, Paul W.","article_content":"During NACPEC expeditions plant species are targeted for collection for a range of reasons including environmental adaptabilities, conservation value, and ornamental features. Presented here are thirteen profiles of notable plants collected on these expeditions. Paperbark Maple Acer griseum Paul W. Meyer P aperbark maple is an iconic Chinese species with beautiful exfoliating cinnamon-colored bark that never fails to grab attention. It is frequently highlighted in public gardens and connoisseurs' gardens throughout the temperate world. It was first introduced to the United States by E. H. Wilson through the Arnold Arboretum in 1907. In addition to its stunning bark, this species is widely admired for its clean, fine-textured foliage, orange-red fall color, and relatively small stature, usually under 35 feet (10.7 meters) tall. It is believed that until recently, all or most paperbark maples in the United States derived from the genetically narrow 1907 Wilson introduction--it consisted of only two plants. Some contemporary seedlings lack vigor, a possible sign of inbreeding over the past century. The re-collection of paperbark maple to introduce greater genetic diversity has been a high priority from the very beginning of NACPEC planning. Of the many hundreds of plants I have observed and collected in China, none were more exciting than finding a grove of wild paperbark maples on Wudang Mountain in Hubei in 1994. Wudang is the site of a famous Taoist temple, and the forests on its slopes have been relatively well protected. On September 21st, 1994, the Hubei expedition team was especially excited to find a scattering of paperbark maples in the understory of a rich, diverse forest at an elevation of 836 meters (2743 feet). It was one of the most biologically diverse habitats that any of us had ever experienced. Dubbed \"horticulture heaven\" by the collectors, we found many choice species including Stewartia sinensis, Cornus kousa, Cornus controversa, Ilex pernyi, and many others growing naturally on this mountainside. The paperbark maples growing on Wudang Mountain were relatively small, growing on a west-facing slope in thin, rocky soil. Being in the understory, most were leggy and the foliage was high off the ground. With careful observation though, we spotted the winged samaras in the upper canopy. Using pole pruners, we were able to collect herbarium specimens and a small seed sample. The following year, in April 1995, NACPEC team members Rick lewandowski, Teicheng Cui, and Ned Garvey spotted an incredible specimen of paperbark maple in the Baxiam Forest Station in Shaanxi, less than 200 kilometers (124 miles) west of Wudang Mountain. They spotted the tree from afar; its leaves had not yet emerged, and they were struck by a distinct warm orange glow reflecting from the tree's bark. The collectors wrote in their journal, \"Holy Hannah! We encountered the biggest specimen we ever saw of Acer griseum. This was incredible... We can't describe the impact of the bark color and the massive stem--the effect was overwhelming.\" This tree measured 81.4 centimeters (32 inches) in diameter at 3 meters (9.8 feet) above the ground. It was estimated to be about 27 meters (88.6 feet) tall. Though no collections of living germplasm resulted, the herbarium specimens, descriptions, and photographic documentation of this individual tree have redefined our thinking about the potential of this species. Paperbark maple is known to frequently produce empty samaras with no viable seeds. That was the case with the Hubei collections. After cutting open many samaras, we found a few (less than 5 percent) with seemingly viable seeds. Fortunately, there were also small seedlings growing under the trees, some of which were Acer griseum 49 RICK lEWANDOWSKI The \"Holy Hannah!\" specimen of paperbark maple spotted during the 1995 expedition to Shaanxi. 50 Arnoldia 68\/2 PAUl MEyER Paperbark maple is noted for its beautiful bark and red-orange autumn foliage color. carefully dug bare-root, stripped of their senescing leaves, and packed in moist sphagnum moss. The seeds never germinated, but these dormant seedlings were brought home successfully and 13 are growing at NACPEC institutions. Though this lack of viability is frustrating to a propagator, the upside is that this species has little potential of becoming invasive. I live on the grounds of the Morris Arboretum and two of the Wudang Mountain paperbark maples are growing in my garden. One is an especially fine specimen. It is a very vigorous individual and in its youth it grew more than 1 meter (3.3 feet) a year. Today it stands over 8 meters (26.2 feet) tall and has a diameter of 27 centimeters (10.6 inches) measured 30 centimeters (11.8 inches) from the ground, just below the first branch. Perhaps because of its strong growth, its bark is especially beautiful, with heavy exfoliation. Morris Arboretum propagator Shelly Dillard took cuttings of this tree in 1998, 1999, 2000, and 2001 while the tree was still juvenile. None rooted successfully. In 2001, I rooted a low branch of the tree in my garden by layering, resulting in the only successful propagation of this individual. This layered plant has the same vigorous habit as its parent, and in July 2010 it had a 6.2 centimeter (2.4 inches) diameter measured 30 centimeters (11.8 inches) from the ground and stood about 3.5 meters (11.5 feet) tall. Four other individuals of this Hubei accession are growing at the Morris Arboretum and each year we watch them, hoping that one might produce some fertile seed for growing on and further evaluation. Bibliography Del Tredici, Peter. 2007. The Paperbark Maple--One Hundred years later. Arnoldia 65(2): 40. Del Tredici et al. 1995. Plant Collecting on Wudang Shan. Arnoldia 55(1): 1220. Paul W. Meyer is the F. Otto Haas Director of the Morris Arboretum of the University of Pennsylvania in Philadelphia, Pennsylvania. "},{"has_event_date":0,"type":"arnoldia","title":"Beautybush Kolkwitzia amabilis","article_sequence":9,"start_page":51,"end_page":52,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25486","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060b326.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Beautybush Kolkwitzia amabilis Michael Dosmann I t was in the late summer of 1901, while exploring the mountains northwest of Ichang, Hubei, China, that Ernest Henry Wilson encountered a shrub which would become one of his favorite introductions: Kolkwitzia amabilis. At the time, he wasn't even quite sure what it was--his notes for collection #1007 simply state that the unnamed plant was 5 feet (1.5 meters) tall, had been free-flowering, with possibly red blooms, and had spinose fruits. The seeds were sent to Veitch Nursery in England where they germinated and grew. In November 1907, plants (labeled as Abelia sp.) were sent to the Arnold Arboretum--the species' first introduction to North America. Shrubs (now under the correct moniker Kolkwitzia) flowered at the Arboretum for the first time in June 1915. Their early-summer displays of pink blossoms, profusely borne on arching branches, so impressed Wilson and others that it was christened beautybush. Thereafter, in early to mid June, the Arboretum's Bulletin of Popular Information routinely included a glowing snippet about the blooming KolMICHAEl DOSMANN Beautybush bears a profusion of pink blossoms in early summer. 52 Arnoldia 68\/2 PETER DEl TREDICI Kevin Conrad stands next to a visually unimpressive but botanically important specimen of beautybush, which was collected from during the 1994 Hubei trip. kwitzia, how big they were getting, and which specimen in the Arboretum was faring best. In fact, the species' merits were lauded to such an extent that in 1927 Wilson noted that the original plant on Bussey Hill had \"been much mutilated for propagation purposes, and from it, either by seeds or cuttings, has originated the whole stock of this plant in America.\" Not everybody agreed with Wilson's endorsement, however, with some even suggesting that plants were not as gorgeous in flower as claimed, or that the plants didn't flower at all. His dander up, Wilson sought to set the record straight on several occasions. His statement on June 7, 1930, (a few months before his untimely death) sums it up: \"There is a foolish rumor abroad that this plant when raised from seed does not blossom. The story is ridiculous since the original plants were raised from seed and the particular plant on Bussey Hill Road is also a seedling... Another canard in circulation is that it is an acid-loving plant. As a matter of fact, it will do equally well on a moderately acid soil or on limestone.\" Nobody could doubt his love for the plant, for in the same passage he states \"Among the deciduous-leaved shrubs that central and western China has given to American gardens Kolkwitzia stands in the front rank.\" Amazingly, it was not until September 25, 1994, that this beautiful and elusive species was collected again, about 100 miles north of Wilson's original collection site in Hubei. The participants on the NACPEC expedition to Wudang Shan discovered multiple plants on a hillside near yan Chi He, and collected ample seed (collector number WD 122). The germinated seedlings at the Arnold Arboretum grew vigorously, and within 18 months were about 0.5 meter (1.6 feet) tall. Unfortunately, all of these plants were sold by mistake at the Arboretum's plant sale in 1997. This was a striking loss, but fortunately seeds of this collection were grown at other institutions also. In the spirit and interest of sharing material, the Arnold Arboretum received cuttings from 3 plants at the Morris Arboretum in 2008. They have rooted and will eventually be planted out. Almost everything we know about this species in cultivation can be traced to Wilson's single introduction event, so we are curious to see how this new collection compares to the original germplasm. No formal studies or evaluations have taken place so far, but there are some preliminary observations that are worth further investigation. Phenological data from the Morton Arboretum from the past 8 years show that the Wilson material on average reaches peak bloom about 1 week earlier than the Wudang Shan material. Perhaps only a minor difference, but this is worth further study. If it holds true, it would be worth selecting for later blooming in self-pollinated F1 and F2 generations of the Wudang Shan germplasm. Michael Dosmann is Curator of living Collections at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Henry's Viburnum Viburnum henryi","article_sequence":10,"start_page":53,"end_page":54,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25494","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed0608928.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Bordelon, Carole","article_content":"Henry's Viburnum Viburnum henryi Carole Bordelon V iburnum henryi, commonly known as Henry's viburnum, was discovered by Augustine Henry and introduced to the west by E. H. Wilson in 1901. Native to central China, V. henryi is relatively rare in cultivation in the United States, existing primarily in public gardens. When this plant was discovered during the fall 1996 NACPEC expedition to the Qinling Mountains in China's Shaanxi province, the team of collectors considered this find a high point of their trip. They were impressed by its beautiful dark green foliage and its large clusters of glossy red fruit. The seeds were collected and propagated for trialing, and fourteen years later, V. henryi is still an impressive plant growing at the United States National Arboretum and the Morris Arboretum. Henry's vibur num is an evergreen shrub, typically growing 7 to 15 feet (2.1 to 4.6 meters) tall, and is hardy in USDA Zones 7 to 10 (average annual minimum temperature 0 to 40F [-17.7 to 4.4C]). Its growth habit is lax, especially when young, but its spreading, arching branches can be pruned into a small upright tree, if desired. Otherwise, little aesthetic pruning is required (any pruning should be done after flowering). The attractive foliage and flowers of Henry's viburnum. DANIEl MOSQUIN, UBC BOTANICAl GARDEN PAUl MEyER 54 Arnoldia 68\/2 DANIEl MOSQUIN, UBC BOTANICAl GARDEN This handsome plant has year-round ornamental interest as well. In the spring months, the new foliage emerges with a bronzy cast that matures to a glossy dark green, which is held throughout the growing season. The narrow, 2 to 5 inch (5 to 13 centimeters) long leaves are serrated above the middle of the leaf and may sport attractive red petioles. During the fall, the leaves may take on a purplish-red hue-- depending on sun exposure--that persists into winter. During winter, the grayish brown bark is revealed. V. henryi blooms in June, displaying slightly fragrant white flowers that occur in panicles that are 2 to 4 inches (5 to 10 centimeters) tall and wide. The flowers are attractive to both bees and butterflies. In July, terminal clusters of glossy red fruit appear, covering the entire plant. As the summer wanes, the red fruit matures to black. Henry's viburnum grows best in full sun or part shade and prefers a well-drained, slightly acid, moist soil. It is not a heavy feeder, but it can be fertilized in late winter and after flowering. Applying a mulch such as composted leaves reduces the need for supplemental watering and fertilizer. No disease or insect problems causing substantial damage to the plant have been noted. The ideal propagation method is by semi-hardwood cuttings some time between late May and late June. Propagation by seed is also possible, but requires at least one cycle of warm\/cold stratification and may take up to several years to germinate. Viburnum henryi is easy to grow and fits into a variety of landscapes. It is recommended for gardens in the southeastern and northwestern sections of the United States but, since it has also performed well in the Washington, D.C., area, it is worth testing in protected sites in Zone 6 areas. Henry's viburnum makes a worthy addition to gardens and may be a more common sight in the future. Carole Bordelon is Supervisory Horticulturist, Gardens Unit, at the United States National Arboretum in Washington, District of Columbia. CAROlE BORDElON Henry's viburnum bears fruit that turns from red to black in late summer. It has an upright, open growth habit. "},{"has_event_date":0,"type":"arnoldia","title":"Manchurian Fir Abies holophylla","article_sequence":11,"start_page":55,"end_page":56,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25496","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070a326.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Meyer, Paul W.","article_content":"Manchurian Fir Abies holophylla Paul W. Meyer F irs are among the most beautiful of landscape conifers. However, since firs are mainly native to cool northern areas or high elevations, many of them do not grow well south of New york City or in regions where summers can be very hot. The Manchurian fir, Abies holophylla, is among the most heat tolerant firs and has proven itself well adapted to midwest and mid-Atlantic states. It is also one of the most handsome firs, with distinctive bright green foliage color and a wide-spreading horizontal branching pattern with age (it is more pyramidal in youth). In addition to its heat tolerance, it is exceptionally winter hardy, capable of withstanding temperatures to -30F (-34C). In nature, the Manchurian fir grows not only in the mountains but also at lower elevations and in valleys where it is exposed to hot summer temperatures. Its range includes North and South Korea, northeastern China, and far southeastern Russia. It often grows in association with Korean pine (Pinus koraiensis), Mongolian oak (Quercus mongolica), purplebloom (or Korean) maple (Acer pseudosieboldianum), and three-flowered maple (A. triflorum). It is among the tallest trees in the forest canopy, often exceeding 100 feet (30.5 meters). In China its wood is valued for use in construction and furniture, though it is not considered as high quality as the wood of Pinus koraiensis. In the northeastern United States, with the decline of eastern hemlock (Tsuga canadensis) and the overuse of white pine (Pinus strobus) and Norway spruce (Picea abies), we have a need for a greater diversity of attractive, welladapted conifers. Since it was first introduced in 1905, Manchurian fir has proven itself to be a useful, non-invasive, and adaptable landscape plant. But unfortunately it is still little known The growth habit and foliage of Manchurian fir. PAUl MEyER 56 Arnoldia 68\/2 PAUl MEyER 25 feet (7.6 meters) tall and 22 feet (6.7 meters) across at the ground. Seedlings from other accessions collected in China in 1993 are just hitting their stride, now standing 4 feet (1.2 meters) tall. At the Morris Arboretum, seeds were treated with cold stratification for 60 days at 41F (5C) before sowing. The seeds then generally germinated reliably within a few weeks. We have found that the seedlings are very slow growing for the first few years, but as they get established can grow over 18 inches (45.7 centimeters) a year. Several specimens of Abies holophylla have been growing well at the Morris Arboretum since before 1933. These mature specimens are over 70 feet (21.3 meters) tall and have taken on a distinctive broadspreading horizontal habit. A prized specimen at the Morton Arboretum in lisle, Illinois, was planted in 1939 and is considered one of their best firs. Curator Kunso Kim reports young plants from the NACPEC expeditions are also performing well at the Morton Arboretum. He observes that the Manchurian fir is relatively This tall specimen of Manchurian fir was photographed in South Korea, part of shade tolerant, although plants its native range. have a more open habit in the shade. Heavy clay or poorly drained soils can outside of arboreta and botanic gardens. Until be problematic for firs, so planting on higher, recently the germplasm represented in this well-drained sites is recommended. country was narrow, not fully representing the landscape-sized Manchurian firs are difficult species' geographic range in its natural habitats. to find in nurseries, but small plants are someThrough the work of collaborative expeditions times listed by specialty mail order nurseries. A to Korea in the 1980s by institutions that later diligent and patient collector will certainly be formed NACPEC, followed by the 1993 and rewarded with a fine growing specimen. 1997 NACPEC expeditions to Hielongjiang and Jilin Provinces, the genetic representation in North America has been expanded. Paul W. Meyer is the F. Otto Haas Director of the Seedlings grown from accessions collected Morris Arboretum of the University of Pennsylvania in Philadelphia, Pennsylvania. in Korea in 1981 are thriving and are now over "},{"has_event_date":0,"type":"arnoldia","title":"Qinling Maple, Acer tsinglingense; or Franchet's maple, Acer sterculiaceum subsp. franchetii","article_sequence":12,"start_page":57,"end_page":58,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25499","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed070ab28.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Justice, Douglas","article_content":"Qinling Maple, Acer tsinglingense; or Franchet's maple, Acer sterculiaceum subsp. franchetii Douglas Justice DOUGlAS JUSTICE C hina serves up an enormous variety of plants for our gardens and landscapes, so much so that the plant explorer Ernest Henry \"Chinese\" Wilson famously called China \"the mother of gardens.\" One need only think of the numbers of cultivated Viburnum, Rhododendron, and Magnolia species that hail from China to get an idea of the magnitude of temperate plant diversity there. While impressive in number, many Chinese species are too tender for cultivation in much of North America. So when a \"new\" cold-hardy Chinese maple comes along, it is cause for gardeners to sit up and take note. Such is the case with a maple collected in the Qinling (Tsingling) Mountains of Gansu Province during the 1996 NACPEC trip: the Qingling maple (Acer tsinglingense)--or Franchet's maple (Acer sterculiaceum subsp. franchetii) as it's being called in North America. Qinling maple is native to the mountains of Shaanxi, Henan, and Gansu provinces in north central China at elevations of 1200 to 1500 meters (3940 to 4920 feet). This area of China is an important bio- By either name, this maple may be a promising addition to North American landscapes. diversity hot spot, with many defined by the Qinling Mountains, where temendemic plant and animal species. Two parallel peratures are considerably cooler than in the mountain ranges--the Qinling and the Daba-- southern Daba Mountains at the same elevatrend east-west, dividing the moist, subtropical tion. According to the collection notes from the to warm temperate south and the drier, cold 1996 expedition, this species was found growtemperate north. The northern boundary is 58 Arnoldia 68\/2 Autumn foliage color of Qinling (or Franchet's) maple. ing on a steep stream bank alongside a variety of familiar temperate plants including Carpinus (hornbeam), Malus (crabapple), and Cornus (dogwood), a good indication that it should be relatively cold hardy. Franchet's maple is found to the southwest of the Qinling Mountains but there is some debate about whether the two species are truly separable. North American botanists generally consider them the same species; however, distinctions have been noted among some individual specimens growing in cultivation. The question is, are the distinctions clear enough to warrant a split (as recommended in Flora of China)? According to the Flora of China account, A. tsinglingense displays three-lobed leaves with wide-spreading side lobes, while the leaves of A. sterculiaceum subsp. franchetii are of a thicker texture and have forward-pointing lobes. The young branches of Qinling maple are described as light brown (vs. darker for Franchet's), and the inflorescences, individual flowers, and samaras are smaller in Qinling maple. In gardens, A. tsinglingense appears to be a robust, medium-sized maple with a strong branch structure that produces an upright-spreading crown. leaves have a papery texture and turn beautiful shades of apricot and red in autumn. It is worth noting that cultivated plants of A. tsinglingense--specimens at the United States National Arboretum and the Morris Arboretum--have often been described as handsome or attractive, while those of Franchet's maple are widely dismissed as dull or coarse. Most European accounts list A. sterculiaceum subsp. franchetii as having little or namental value, and plants at the University of British Columbia Botanical Garden grown from older seed collections made in Hubei and Sichuan Provinces (to the south and west of the Qinling collections) could also easily be described in this disapproving light. On the other hand, the response of maples to the climate in eastern North America is often manifested in neater, more compact growth and autumn leaves with more saturated colors. This could explain much of the difference, but until a wider sample--representing trees from the Qinling and beyond--are grown under the same conditions, these questions will go unanswered. Whether we are seeing a minor variant of Franchet's maple or a bona fide species in Qinling maple is an open question. More research is required to settle the science, but judging by the plants in gardens, this fine-looking maple appears worthy of wider cultivation, at least in eastern North America. Douglas Justice is Associate Director and Curator of Collections at the University of British Columbia Botanical Garden in Vancouver, British Columbia. DOUGlAS JUSTICE "},{"has_event_date":0,"type":"arnoldia","title":"Chinese Stewartia Stewartia sinensis","article_sequence":13,"start_page":59,"end_page":60,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25491","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed0608126.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Peter Del Tredici I n perusing my handwritten journal from the 1994 NACPEC trip to Wudang Shan in Hubei Province, China, I found the following entries regarding the Chinese stewartia, Stewartia sinensis: Monday, September 19, 1994: \"The other highlight [besides finding Acer griseum] was to see Stewartia sinensis--the biggest plant I've seen of any Stewartia--about 30 inches [76 centimeters] in diameter at the base and 50 feet [15 meters] tall, with a clear bole for the first 20 to 30 feet [6 to 9 meters]. The bark was a buff\/tan color-- carried high up into the crown--and absolutely smooth with little or no flaking. Certainly the most magnificent tree I saw on Wudang Shan. It should also be noted that a spot right next to the Stewartia, above it to be precise, was selected as a site for a public toilet--and it was disgusting to the point that no one but me dared to go near the tree. Perhaps that is the secret to its vigor.\" Tuesday, September 20, 1994: \"Up the mountain again, then a quick turn to the east at about 1300 meters [4,265 feet] and off into the woods. It was misty and rainy the whole day, giving the whole place a great air of mystery... The rain and heavy This young specimen of Chinese stewartia displays multi-colored bark. fog limited our visibility pretty much to what was immediately beside the path--but kousa was there too, and a small (4 inch [10 centhere was so much. After about an hour or so timeter] diameter) specimen of Acer griseum. All on the path, we came upon an incredible house within the narrow space of ten square meters--I nestled under the cliffs. It looked like it had been felt as though I'd died and gone to horticultural there forever--no one was around so I took a few heaven. The only thing missing, sadly, was seeds pictures. It really felt like the same China that on any of the plants. The conditions were moist Wilson saw. A little ways beyond the house and and shady and steep, with an oak overstory.\" garden we came to a bend in the road where it It was in this location--Hubei Horticultural looped back into a ravine. Our guide, Mr. Zeng, Heaven we called it--that I noticed a small a collector of medicinal plants among other Stewartia seedling, about 20 centimeters (8 things, pointed out a beautiful specimen of Stewinches) tall with a distinct kink at its base, artia sinensis, and then we saw another, both growing along the edge of the path. I immewith the beautiful, rich, smooth, cinnamon-red bark--a wonder to behold and to touch! Cornus diately stopped and dug it up while the other PETER DEl TREDICI Chinese Stewartia Stewartia sinensis 60 Arnoldia 68\/2 PETER DEl TREDICI Chinese stewartia blooms in midsummer. PETER DEl TREDICI members of the party went on ahead. At the end of the expedition, the plant was washed clean of soil and packed in moist sphagnum moss for shipment back to the United States. The plant passed inspection at the USDA inspection station at Beltsville, Maryland, and, after 48 days on the road, arrived at the Arnold Arboretum on November 7, 1994. It was immediately potted up, assigned the accession number 691-94, and left to overwinter in a cool greenhouse. The seedling was moved to the outdoor nursery in spring 1996, and it grew to a height of 1.1 meters (3.6 feet) by the end of July. In spring 2000, it was moved from the nursery to the grounds, where it has proved to be completely hardy. By the end of the 2009 growing season, the plant was 4.7 meters (15.4 feet) tall and 4.1 meters (13.5 feet) wide with two co-dominant trunks arising from the stout base which is 14 centimeters (5.5 inches) in diameter. The plant flowered for the first time in the summer of 2002 and has gone on to flower every year since. The flowers, which are produced from the end of June through mid July, are 6 to 7 centimeters (2.4 to 2.8 inches) across with beautiful light lemon yellow anther filaments and short pedicels, indicating that it is S. sinensis var. sinensis according to the Flora of China. In Boston, the new leaves emerge in early May and are distinctly hairy and tinged with red anthocyanin pigment; both features disappear within a week or so. The bark is cinnamon-red where it is not exfoliating and olive-green where the bark has peeled off in fine, papery flakes. Over time, I'm sure the tree will develop mature bark that is \"smooth as alabaster and the colour of weathered sandstone\" (to quote W. J. Bean), like the magnificent specimen we saw on Wudang Shan. Peter Del Tredici is a Senior Research Scientist at the Arnold Arboretum. A mature specimen in China shows smooth, sandstone-colored bark. "},{"has_event_date":0,"type":"arnoldia","title":"Chinese Chinquapin Castanea henryi","article_sequence":14,"start_page":61,"end_page":62,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25489","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060bb28.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Anagnostakis, Sandra L.","article_content":"Sandra L. Anagnostakis C harles Sprague Sargent wrote a beautiful description of this chestnut tree in his Plantae Wilsonianae in 1917. He also revised the taxonomy to the name that we still use today. Here is an excerpt from his article: \"This very distinct species is distributed from the neighborhood of Ningpo through the valley of the Yangtsze River as far west as Mt. Omei. On the mountains of western Hupeh and of eastern Szech'uan it is common in woods. This chestnut grows to a larger size than any other Chinese species and trees from 20 to 25 meters [66 to 82 feet] tall with trunks from 1 to 3 meters [3.3 to 9.8 feet] are common. Occasionally trees 30 meters [98.4 feet] tall and 5 meters [16.4 feet] in girth of trunk are met with. The leaves are green on both surfaces and entirely glabrous except for a few appressed hairs on the underside of the primary and secondary veins. The leaves are without lepidote glands except on the upper surface of the very young leaves, E. H. Wilson made this photograph of a large Castanea henryi growing in from which they disappear very early. Hubei Sheng, China, on June 22, 1910. Although variable in size the leaves are very characteristic; they are always caudatelaway was very interested in chestnuts, and acuminate and broadest below or at the middle, in 1935 he began planting chestnut trees from and the secondary veins are projected in long Asia procured by the United States Department aristate points. The shoots are dark-colored and of Agriculture's plant exploration and imporquite glabrous and the winter-buds are brownish, tation program. Over a period of eight years, short, broadly ovoid, obtuse or subacute and are he planted 2,192 chestnuts, and among them glabrous or nearly so. The styles vary in number were 202 Castanea henryi from eight differfrom 6 to 9, and the fruit may be solitary or two ent locations in China. I visited the chestnut or three on a short spike. The spines of the ripe involucre are sparsely villose. All the fruits we plantings at Callaway Gardens in 1993 with have seen contain a solitary nut, but it is probDr. Jerry Payne and Ann Amis from the USDA. able that occasionally two occur, as they do in We noticed that most of the chestnuts were the American C. pumila.\" badly damaged by the Asian chestnut gall wasp My interest in this species was piqued when I saw a large planting of it at Callaway Gardens in Hamilton, Georgia. Founder Cason J. Cal- that Dr. Payne had discovered and described in 1976, shortly after a chestnut grower accidentally brought it to central Georgia. Within one ARCHIveS OF THe ARNOlD ARbOReTUM Chinese Chinquapin Castanea henryi 62 Arnoldia 68\/2 PAUl MeYeR This Morris Arboretum specimen of Castanea henryi (WD-069) was grown from seed collected in Wudang Shan on the 1994 Hubei expedition. It flowers heavily and bears sweet-tasting nuts that are quickly devoured by squirrels, deer, and other wildlife. PAUl MeYeR C. henryi, and all plants that we saw of this species had either very few or no galls. Finding a chestnut species that was clearly resistant to gall wasp presented an opportunity to breed resistance into our orchard and timber chestnut lines (C. henryi is also resistant to chestnut blight). We have one mature C. henryi here at the Connecticut Agricultural experiment Station, but others planted over the years have not been winterhardy enough to survive. Seeds that Wilson collected were planted at the Arnold Arboretum, but no trees from this accession (AA-551) now survive. However, one open-pollinated offspring (a probable cross with a nearby Chinese chestnut, C. mollissima) is still alive and well ( AA-623-32). Since that discovery of gall wasp resistance in Georgia, I have been including C. henryi in my crosses to produce better timber and nut-producing chestnut trees for our northeastern forests and orchards. We don't know what pest or disease of chestnuts will next be brought into the United States, but it is clear that imported chestnut species will be called into use in hybridization to combat these new threats. All the more reason to say \"Keep exploring, NACPeC!\" References: Payne, J. A., R. A. Green, and C. D. lester, 1976. New nut pest: an Oriental chestnut gall wasp in North America. Annual Report of the Northern Nut Growers Association. 67: 8386. Sargent, C. S. 1917. Plantae Wilsonianae: An enumeration of the woody plants collected in western China for the Arnold Arboretum of Harvard University during the years 1907, 1908, and 1910 by E. H. Wilson. volume 3, p. 197. Cambridge, Massachusetts: University Press. Developing nuts are enclosed in prickly involucres. of the plantings we came to a block of what I thought were Japanese chestnuts (C. crenata) and noticed that there were no galls. A check of the planting plan revealed that these were Sandra l. Anagnostakis is a research scientist in plant pathology and ecology at the Connecticut Agricultural experiment Station in New Haven, Connecticut. eRRATUM 15 December 2010: The specimen shown in the two images above has been identified as a Castanea hybrid rather than straight species Castanea henryi. "},{"has_event_date":0,"type":"arnoldia","title":"Epimediums Epimedium spp.","article_sequence":15,"start_page":63,"end_page":64,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25492","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060816b.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Bordelon, Carole","article_content":"Carole Bordelon A lthough the vast majority of the plants targeted for collecting by NACPEC are trees and shrubs, several genera of herbaceous plants have been targeted as well. Since China is the major area for the diversity of epimediums (Epimedium spp.), it made perfect sense to target this group of adaptable perennials. Thirteen accessions of epimedium were collected during Epimedium stellulatum bears starry white flowers. NACPEC trips, several of which are highly ornamental and should be recognized as worthy plants for the shade garden. In the fall of 1994, members of the NACPEC expedition to the Wudang Shan mountain range located in central China (which is famous for its exceptionally rich and diverse flora) collected several noteworthy accessions of epimedium. This was a historically significant trip as there were many new species of epimedium being described in China at the same time. With the help of Darrell Probst, an expert in the collection, identification, and introduction of epimediums, those 1994 Wudang Shan epimedium accessions were correctly identified and, more importantly, several of them proved Epimedium lishihchenii has long-spurred yellow flowers. to be new species not represented in the NACPEC members' institutional holdings. -34.4C]). It has a running habit and reaches 12 Epimedium lishihchenii and Epimedium stelinches (30.5 centimeters) in height. The large lulatum are just two of the species identified evergreen leaflets are leathery and sustain little from that trip. damage in the winter months. It has bright yelEpimedium lishihchenii is an attractive plant low flowers with elongated spurs, and blooms hardy to at least USDA Zone 4 (average annual in late April or early May. Although the flowers minimum temperature -20 to -30F [-28.9 to are attractive, this plant's best quality is the CAROlE BORDElON CAROlE BORDElON Epimediums Epimedium spp. 64 Arnoldia 68\/2 foliage. It remains fairly clean throughout the growing season. In the winter, the foliage may be tinged with an attractive burgundy color. like Epimedium lishihchenii, Epimedium stellulatum is an evergreen species hardy to Zone 4. It has a clumping habit and is smaller in stature than Epimedium lishihchenii. E. stellulatum blooms early in the season, revealing small white starlike flowers that are held above the leaves on erect stems. Though it is considered to be evergreen, it has sustained more winter damage to the foliage at the United States National Arboretum than Epimedium lishihchenii. During the 1996 NACPEC expedition to the Quinling Mountains, several additional accessions of Epimedium stellulatum were collected. These additional accessions have leaflets that are slightly larger and narrower than the 1994 Epimedium stellulatum collection. CAROlE BORDElON The epimediums that hail from China occur primarily in woodlands in temperate hilly or montane regions. Keeping this in mind, they do best in moderately cool and half shady conditions. They thrive best in a moist but well drained soil and will tolerate periods of drought provided their roots are not exposed. They benefit from being mulched with leaf mold. Epimediums are best propagated by division. Epimediums make excellent landscape plants because they can grow in a variety of situations and are easy to care for. Depending on the species, they thrive in sun or full shade and can provide year-round interest in gardens. Carole Bordelon is Supervisory Horticulturist, Gardens Unit, at the United States National Arboretum in Washington, District of Columbia. Glossy, leathery foliage is an ornamental feature of Epimedium lishihchenii. "},{"has_event_date":0,"type":"arnoldia","title":"Chinese Hemlock Tsuga chinesis","article_sequence":16,"start_page":65,"end_page":67,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25490","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060bb6d.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Chinese Hemlock Tsuga chinensis Peter Del Tredici A specimen of Tsuga chinensis var. tchekiangensis growing in Jiangxi. RiCk LEWANdOWSki O ne of the most important of all NACPEC collections is the Chinese hemlock (Tsuga chinensis). Prior to 1979, this species appears to have been successfully introduced into North America only once--a single seedling collected by E. H. Wilson in Hubei, China, in 1910 that is still alive today. This accession has been frequently propagated and widely distributed by the Arnold Arboretum. The lack of Chinese hemlock diversity became a significant factor when horticulturists began to notice that the species was highly resistant to the Japanese strain of hemlock woolly adelgid (HWA) that was ravaging native stands of eastern hemlock (Tsuga canadensis) throughout the central and southern portion of its range in eastern North America. Working through its various Chinese contacts, NACPEC began a concerted effort to acquire Chinese hemlock germplasm in order to facilitate research on its growth rate, habitat tolerances, and resistance to HWA. The collections began in 1994 with seeds provided by the Xian Botanical Garden and peaked in 1996 with 6 separate collections from various habitats in the Qinling Mountains in Shaanxi Province, the northern part of 66 Arnoldia 68\/2 PETER dEL TREdiCi Tsuga chinensis growing in montane habitat. its range. in all, some 33 different collections of three different varieties of the species were made. Representatives from 19 of these collections--totaling some 250 plants--are growing at various NACPEC gardens. The largest of them is at the Morris Arboretum and has reached 4.9 meters (16 feet) in height with dBH (diameter at breast height) of 9 centimeters (3.5 inches) after 10 years of growth. True to initial reports, Chinese hemlock has so far proved completely resistant to HWA in a wide variety of North American locations. it is fully cold hardy into USdA Zone 5 (average annual minimum temperature -10 to -20F [-23.3 to -28.9C]) and is relatively fast growing--the mean height of 38 seedlings growing under variable conditions on Hemlock Hill at the Arnold Arboretum was 169 centimeters (66.5 inches) at 10 years of age. i observed Chinese hemlock at the Arnold Arboretum during the 2009 growing season and noted that the species both began growing and stopped growing about two weeks earlier than eastern hemlock. interestingly, the new growth on vigorous terminal shoots was the same for both species--about 45 centimeters (17.7 inches)--which was the greatest among the seven hemlock species measured. Compared to eastern hemlock, Chinese hemlock is a bit Tsuga chinensis 67 PETER dEL TREdiCi Slightly drooping branch habit on a young Chinese hemlock. Attractive foliage of Chinese hemlock. woolly adelgid (Adelges tsugae). Journal of Arboriculture 30(5): 282287. Bentz, S. E., L. G. H. Riedel, M. R. Pooler, and A. M. Townsend. 2002. Hybridization and selfcompatibility in controlled pollinations of eastern North American and Asian hemlock (Tsuga) species. Journal of Arboriculture 28(4): 200205. Hooper, B. k., R. M. Bates, J. C. Sellmer, and G. A. Hoover. 2009. Challenging Chinese hemlock (Tsuga chinensis) with hemlock woolly adelgid (Adelges tsugae) ovisacs. Arboriculture and Urban Forestry 35(1): 14. Peter del Tredici is a Senior Research Scientist at the Arnold Arboretum. \"droopier\" during the growing season, seems to be comparably shade tolerant (although this trait has not actually been quantified), is less cold hardy--Zone 5 versus Zone 3 (average annual minimum temperature -30 to -40F [-34.4 to -40C])--and, as mentioned, has the great advantage of adelgid resistance. it is a terrific plant for replacing adelgid-killed eastern hemlocks under cultivated conditions. References del Tredici, P. and A. kitajima. 2004. introduction and cultivation of Chinese hemlock (Tsuga chinensis) and its resistance to hemlock PAUL MEyER "},{"has_event_date":0,"type":"arnoldia","title":"Amur Maackia Maackia amurensis","article_sequence":17,"start_page":68,"end_page":70,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25485","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060af6d.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Meyer, Paul W.","article_content":"Amur Maackia Maackia amurensis Paul W. Meyer O ne of our most impor tant plant exploration goals is collecting tree species that demonstrate stress tolerance and are therefore likely candidates for evaluation as urban street trees. Maackia amurensis is outstanding in its promise as a tough and useful urban tree. It is native over a wide geographic area including Japan, the Korean peninsula, northeast China, and far eastern Russia. It is a member of the legume family (Fabaceae) and it is one of the relatively few trees that support nitrogen fixing bacteria on its roots. Although it was introduced to the United States in the late nineteenth century, it is still relatively rare here. In cultivation Maackia amurensis is a mediumsized tree reaching 45 feet (13.7 meters) or more (we were surprised to see specimens in China growing well up into the forest canopy, taller than the species is usually reported to grow). It has compound leaves similar to its relative, black locust (Robinia pseudoacacia). As the leaves emerge in the spring they are covered with silky hairs which give the tree a silvery-gray appearance. A 40-foot tall specimen of Amur maackia at the Morris Arboretum. PAUl MEyER Maackia amurensis 69 PAUl MEyER This Amur maackia in China managed to survive in difficult soil conditions with highly variable moisture levels, an indicator that the species may perform well as an urban street tree. PAUl MEyER Newly emerged foliage of Amur maackia has a silvery sheen. Upright racemes of small creamcolored flowers appear in July, a time when few other trees are blooming. Amur maackia's bark is slightly exfoliating with handsome shades of copper and tan. It is especially striking when backlit. Professor Jin Tieshan of the Heilongjiang Academy for Forestry reported that Amur maackia's darkcolored wood is very valuable and in the 1990s it was commonly exported to Japan. He also pointed out that on twigs the young sapwood is a light greenish tan, while the older heartwood takes on a dark brown color. The wood is exceptionally hard and rot resistant; traditionally, it has been used for fencing in China, similar to the use of black locust wood in the United States. NACPEC explorers collected 3 accessions of this species in 1993 in Heilongjiang. Collection HlJ085 was made along Jiang Po lake, where it grew along the high water line in thin, sandy soils overlaying rock. It was clear that the trees had to tolerate wet soils when the water levels were high and then very droughty conditions when the water level dropped. Adaptability to these kinds of natural conditions suggests that this species might also be adapted to the periodic root flooding and droughts that plague urban street trees. At another site in Heilongjiang, small, stunted Amur maackias were growing on a rocky, ancient lava flow along a stream edge. This area was subject to alternating periods of flooding and drought. Few other woody plants could survive there, but these trees were able to withstand the difficult environment, growing out of fissures in the rock. We tried two different treatments to soften the hard outer coats of Amur maackia seeds from collection 70 Arnoldia 68\/2 Amur maackia bears upright racemes of creamy white flowers in midsummer. PAUl MEyER HlJ051. One group of seeds was given a 24-hour soak in hot water and a second group of seeds was scarified in sulfuric acid before being sown. Both treatments resulted in excellent germination. Three 16-year-old specimens are growing near my home on the grounds of the Morris Arboretum. These Amur maackias stand 18 feet (5.5 meters) tall and are 5 inches (12.7 centimeters) DBH (diameter at breast height). They have attractive foliage, flowers, and bark, and are handsome in every season of the year. They are growing on a hot, sunny, south-facing slope and have never exhibited stress in times of drought. Maackia amurensis is certainly proving itself to be a handsome, tough, adaptable shade tree that should be used more widely in stressful urban sites. Bibliography Pai, J. G. B. and W. R. Graves. 1995. Seed source affects seedling development and nitrogen fixation of Maackia amurensis. Journal of Environmental Horticulture 13:142146. Paul W. Meyer is the F. Otto Haas Director of the Morris Arboretum of the University of Pennsylvania in Philadelphia, Pennsylvania. PAUl MEyER Clean green foliage and exfoliating bark add ornamental appeal to Amur maackia. "},{"has_event_date":0,"type":"arnoldia","title":"Farges Filbert Corylus fargesii","article_sequence":18,"start_page":71,"end_page":72,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25493","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060856f.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Aiello, Anthony S.","article_content":"Farges Filbert Corylus fargesii Anthony S. Aiello ANTHONy AIEllO O ne of the goals of plant exploration is to introduce new species into cultivation, and it is a rare and exciting opportunity for any plant collector to do so. Corylus fargesii (Farges filbert) was first described in China by Western botanists in the late 1800s and early 1900s. Although herbarium specimens were collected during this \"golden age\" of plant exploration, there is no evidence that living specimens were grown in arboreta and botanic gardens from these early collections. The first NACPEC collection of C. fargesii occurred on the 1996 expedition to Shaanxi and Gansu, and it was collected again on the 2005 expedition to Gansu (identified by collector numbers QlG-231 and NACPEC05-047, respectively). Until the introduction of Farges filbert seeds to the United States in 1996, little was known or written about The attractive pyramidal form of Farges filbert. this species. ern Ningxia, Shaanxi, and northeast Sichuan Of the many taxa collected by NACPEC over (Chengkou Xian). During the 1996 expedition, the past twenty years, few arouse more exciteFarges filbert was collected in eastern Gansu at ment than Corylus fargesii. The trees display the Xiao long Shan Forest Bureau, Dang Chuan exfoliating tan and copper bark that rivals the Forest Station. In their field notes, the collecmost attractive birches and is especially remitors described it as a truly beautiful tree with niscent of river birch, Betula nigra. According exquisite bark. The parent trees had reached to the Flora of China, Farges filbert grows to 12 to 15 meters (39 to 49 feet) tall and were 40 meters (131 feet) tall and occurs naturally in found growing among rocks in sandy silt loam mountain valley forests at elevations from 800 soil approximately 2 meters (6.6 feet) above a to 3000 meters (2,625 to 9,843 feet) in southern stream in open woodland. Gansu, Guizhou, Henan, Hubei, Jiangxi, south- 72 Arnoldia 68\/2 ANTHONy AIEllO Farges filbert displays exfoliating bark similar to that of river birch. 10 plants from this accession, all of which show remarkably similar growth habit and size. These trees have grown quickly, reaching 25 to 30 feet (8 to 9 meters) after 13 years, with strong central leaders and very uniform broadly ovate habits. The trees exhibit some variation in the level of exfoliation and color of the bark, which ranges from a deep copper to a pale cream color. Farges filbert has clean summer foliage with no insect or disease problems, and turns a good yellow in autumn. Our plants are growing in several locations, with slight differences in soil pH and all with evenly moist, well-drained soils. Farges filbert has been propagated successfully from seeds and by grafting, but with only marginal success from stem cuttings. The diversity of conditions under which it is growing successfully indicates broad adaptability from the central Midwest to New England and south to the mid-Atlantic states. The species appears to be fully cold hardy in USDA Zones 5 through 7 (average annual minimum temperatures -20 to 0F [-29 to -18C]). This tree's highly ornamental exfoliating bark and rapid growth rate indicate great potential as an ornamental tree for a range of situations, and it promises to be an excellent addition to landscapes in the future. Literature Cited Aiello, A.S. 2006. Plant collecting on the eaves of the world. The Plantsman 5(4): 220225. Aiello, A.S. and S. Dillard. 2007. Corylus fargesii: A New and Promising Introduction from China. Proc. Intl. Plant Prop. Soc. 57: 139143. Grimshaw, J. and R. Bayton. 2009. New trees: recent introductions to cultivation. Royal Botanic Gardens, Kew, England. Wu, Zheng-yi and Peter H. Raven, eds. 1994. Flora of China. Missouri Botanical Garden Press. St. louis. On the 2005 NACPEC expedition to southern Gansu, on a morning when we shared our hike through pastures and woodland with numerous cattle, we were fortunate to encounter and collect C. fargesii in Zhou Qu county, near the Sha Tan Forest Station. The parent plants were growing in a mesic mixed deciduous forest and were located a few meters above a stream. Many C. fargesii were seen throughout a small area; all of these trees had been heavily coppiced but had resprouted vigorously. As in 1996, we were impressed by the beautiful exfoliating bark. Corylus fargesii is now well established at all of the NACPEC member gardens and in several other North American public gardens. Trees from the 1996 collection are doing especially well at the Morris Arboretum, where we have Anthony S. Aiello is the Gayle E. Maloney Director of Horticulture and Curator at the Morris Arboretum of the University of Pennsylvania in Philadelphia, Pennsylvania. "},{"has_event_date":0,"type":"arnoldia","title":"Chinese Ashes Fraxinus spp.","article_sequence":19,"start_page":73,"end_page":74,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25488","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060b76f.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Bachtell, Kris R.; Siegel, Olivia","article_content":"Chinese Ashes Fraxinus spp. Kris R. Bachtell and Olivia Siegel S ince its discovery in Detroit, Michigan, During previous NACPEC expeditions, colin 2002, the emerald ash borer (EAB), lecting Fraxinus species had not been a primary Agrilus planipennis, is estimated to have focus. Typically, there were too many other killed over 40 million ash trees in southeastern more exciting species to collect and there was Canada and in 14 states throughout the central no reason to focus on ash. With the invasion of and eastern United States. Native to Asia, EAB EAB this changed. A 2006 NACPEC-sponsored probably arrived in North America by ship in grant request was funded by the USDA and supsolid wood materials used for packing freight. ported contract collecting of native Fraxinus This insect's impact has cost municipalities, seeds in China by Kang Wang of the Beijing property owners, nursery operators, and the forBotanic Garden from 2007 through 2010. Addiest products industry tens of millions of dollars. It appears that no North American ash species is resistant to this pest, so all are threatened--there are an estimated 8 billion ash trees currently growing in the United States. On the few ash species that have been studied in China, EAB is usually a secondary or periodic pest, infesting only stressed trees and not necessarily resulting in tree mortality. Most outbreaks in China have been associated with urban and restoration plantings involving North American species, particularly green ash (Fraxinus pennsylvanica) and velvet ash (F. velutina), both of which have been extensively planted in many northern Chinese cities. It is extremely important to gain access to Fraxinus species from China to test the full range of EAB response and to assess the adaptation of Asian ash species to American conditions and their appropriateness for urban landscapes. There are 22 Fraxinus species listed in the Flora of China. Some of these species are tropical, and therefore not suitable for regions of the United States currently under siege by EAB, but they are of potential utility for expanding the range of ash adaptation or for responding to EAB if the pest proves to be adapted to tropical or subtropical areas Collecting seeds of Manchurian ash (Fraxinus mandshurica) in China. in the New World. KRIS BACHTEll 74 Arnoldia 68\/2 KRIS BACHTEll tionally, the 2008 NACPEC expedition to Shaanxi Province focused on collecting Fraxinus in the botanically rich Qinling mountain region. We collected several thousand seeds of five Fraxinus species there. Several of these species are poorly represented in the United States; for example, Pax's ash (F. paxiana) and island ash (F. insularis) are being grown at only two or three botanical institutions. We made several collections of these species, along with Chinese ash (F. chinensis), Manchurian ash (F. mandshurica), and Chinese flower- Fraxinus insularis foliage and seeds. ing ash (F. stylosa). Of these, ing efforts, noting that the previous lack of Manchurian ash is probably the best known, Asian ash accessions in the United States had since it is a large-growing tree with an estabgreatly restricted the potential for hybridizalished landscape value (the cultivated selection tion. Future collecting expeditions are planned `Mancana' is common in the nursery industry). and many additional research questions related Other species, such as island ash and Chinese to EAB still need to be answered. flowering ash, are medium-sized trees that may have urban use potential if they prove adaptBibliography able. Pax's ash is a shrubby species that possesses extremely large flower clusters, but its Flora of China. (1996) Retrieved December 15, 2006 from http:\/\/flora.huh.harvard.edu\/china\/mss\/ landscape potential is unknown. volume15\/Oleaceae.published.pdf Twenty-six Fraxinus seed collections have Koch, J. l., D. W. Carey, M. E. Mason, and M. N. been made in China, resulting in 11 different Islam-Faridi. (online publication in process). taxa currently represented by over 600 plants. Overcoming obstacles to interspecies Ash seed is relatively easy to harvest, clean, hybridization of ash. In: Proceedings of and germinate, so the degree of success growthe Symposium on Ash in North America, ing these plants has been good. Surplus seeds March 1921, 2010. Purdue University, West lafayette, IN. will be preserved in the USDA's germplasm repository in Ames, Iowa, and made available liu, H., Bauer, l.S., Gao, R., Zhao, T., Petrice, T.R., and Haack, R.A. (2003). Exploratory for scientific research throughout the world. survey for the Emerald Ash Borer, Agrilus Currently, plants from the 2008 Shaanxi expeplanipennis (Coleoptera: Buprestidae), and its dition are being used by leading researchers in natural enemies in China. The Great Lakes efforts to identify relative susceptibility and Entomologist, 36(3 & 4): 191204. resistance of different ash species to EAB. The Poland, T.M., & McCullough, D.G. (2006). Emerald ash seed collections will also aid research in idenborer: Invasion of the urban forest and the threat tifying appropriate genetic material to create to North America's ash resource. Journal of Forestry, April\/May 2006: 118123. new North American-Asian ash hybrids that combine resistance genes from the Asian species with useful characteristics from the North Kris R. Bachtell is Vice President of Collections and American species. Researchers Koch et al. Facilities at the Morton Arboretum in lisle, Illinois. recently cited the importance of the NACPEC Olivia Siegel is Development Coordinator at the Aspen Center for Environmental Studies in Aspen, Colorado. Fraxinus collections to their genetics and breed- "},{"has_event_date":0,"type":"arnoldia","title":"Manchurian Catalpa Catalpa bungei","article_sequence":20,"start_page":75,"end_page":76,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25495","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060896d.jpg","volume":68,"issue_number":2,"year":2010,"series":null,"season":null,"authors":"Olsen, Richard T.; Kirkbride Jr., Joseph H.","article_content":"Manchurian Catalpa Catalpa bungei Richard T. Olsen and Joseph H. Kirkbride, Jr. F amiliarity breeds contempt--for catalpa. The genus has nine accepted species, two in eastern North America, four in the West Indies, and three more in eastern Asia. The North American species (Catalpa bignonioides and C. speciosa) are the best known in the West, often overlooked as waifs in urban landscapes or as country trees, too large or messy for modern landscapes, yet tolerated when in flower in late May to June. Perhaps this sentiment permeated the group conscience on NACPEC's 1994 Wudang Shan expedition, when, on the first day of collecting, they were underwhelmed by Kevin Conrad's sighting of a lone pollarded catalpa in a field of soybeans and corn. Conrad, representing the United States National Arboretum (USNA), was on his first expedition to China, the wide-eyed and energetic newbie in a field of veteran collectors. Reluctantly, the group stopped to collect seed and vouchers of what was identified as the Manchurian catalpa, Catalpa bungei. As it turned out, it was the only catalpa seen on the trip and proved to be one of the most important collections of the expedition. In 1831, Alexander Andrejewitsch von Bunge collected herbarium specimens of a catalpa near Beijing, which C. A. Meyer later identified and described as a new species, and named in honor of Bunge. Catalpa bungei and C. ovata are the two most commonly cultivated species of catalpa in China, both in agroforestry for their high quality wood and in religious circles as one of the \"jeweled trees\" of Chinese Buddhism. There is much research published in Chinese journals on propagation, breeding behavior, and sylviculture of C. bungei, but this emphasis on its importance has not filtered to the West, where appreciation for the species is lacking. Early introductions of catalpa into Europe were erroneously ascribed to this new species, even as they came into flower with upright, many- flowered panicles of small yellow or yellowishgreen flowers that clearly identified them as C. ovata. The inflorescences of C. bungei are corymbose, with fewer but larger flowers conspicuously spotted with pink, effectively coloring the flowers rose. The first introduction of true C. bungei is attributed to the Arnold Arboretum in 1904, when wild-collected seeds were acquired (via American diplomat E. T. Williams) from the vicinity of Beijing. These seeds--and subsequent plants--were distributed to European botanical institutes, but the species remains almost nonexistent in cultivation, a victim of confusion generated by earlier misidentifications. The Arnold Arboretum still has a living plant of this accession (AA#12927), which has stood sentinel above the lilac collection for over a century. The great plant collectors E. H. Wilson and F. N. Meyer did not overlook catalpa on their forays across China in the early twentieth century. Wilson, collecting for the Arnold Arboretum, never knowingly collected C. bungei, but based on herbarium specimens from his trips, his collections of C. fargesii (syn. C. duclouxii) are a mixed bag of phenotypes, some of which agree with C. bungei. But Meyer, collecting for the USDA, collected what he labeled C. bungei on five separate occasions, calling the species \"one of the finest flowering trees in the world\". The taxonomy of these species is not well resolved, but based on recent phylogenetic analyses, this group forms a clade separate from the North American and West Indian species. The USNA conducts on-going taxonomy and breeding work in the genus Catalpa, and C. bungei has taken center stage, thanks to its beautiful flowers, disease resistant foliage, and general adaptability. In our search for germplasm to introduce into our breeding program, we have scoured both domestic and foreign nurseries for material of C. bungei and related species. 76 Arnoldia 68\/2 We have yet to find a nursery offering the real C. bungei. Our search of botanical gardens and arboreta yielded only three accessions in North America that are true-to-type, two of wild origin: the original C. bungei 12927 at the Arnold, and C. bungei WD009 from the Wudang Shan trip in 1994. Unknowingly, NACPEC had made the first collection of C. bungei in 90 years, proHARVARD UNIVERSITy HERBARIA viding germplasm for urban tree breeding and increasing our knowledge of an underutilized and underappreciated genus. Richard T. Olsen is a Research Geneticist and Joseph H. Kirkbride, Jr. is a Plant Taxonomist, both at the United States National Arboretum in Washington, District of Columbia. An herbarium specimen from the lone Catalpa bungei collected in Wudang Shan during the 1994 NACPEC expedition to Hubei, showing its foliage and long seed pods (bent to fit on sheet). Facing inside back cover: Manchurian catalpa (Catalpa bungei) bears lovely rose-tinted flowers. Photo by Richard Olsen. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23421","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160b36e.jpg","title":"2010-68-2","volume":68,"issue_number":2,"year":2010,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Magnolias at the Scott Arboretum of Swarthmore College","article_sequence":1,"start_page":2,"end_page":12,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25484","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060af28.jpg","volume":68,"issue_number":1,"year":2010,"series":null,"season":null,"authors":"Bunting, Andrew","article_content":"Magnolias at the Scott Arboretum of Swarthmore College Andrew Bunting F SCott ARBoREtum rom the inception in 1929 of the Scott Arboretum of Swarthmore College, the mission has remained the same--to collect and display outstanding ornamental plants, specifically trees, shrubs, and vines. Since 1931, one of our most prominent collections of plants--and one that has stood the test of time--has been the magnolia collection. Early on, new magnolia accessions were received from notable nurseries, organizations, and individuals including Bobbink and Atkins, Rutherford, New Jersey; Andorra Nursery, Chestnut Hill, Part of the magnolia collection at the Scott Arboretum. Magnolias at the Scott Arboretum 3 SCott ARBoREtum Pennsylvania; the Arnold Arboretum; Hicks Nursery, Long Island, New York; and Highland Park, Rochester, New York. At the time, John Wister, first director of the Scott Arboretum, was developing the campus based on an evolutionary or phylogenetic tree, so all genera in a plant family were planted together, and hence all species in a family resided together. the magnolia collection housed both species and cultivars alike. In 1931, Wister began to get regular deliveries of many plants, especially magnolias, from Henry Hicks of Hicks Nursery on Long Island, New York. on may 8th, 1934, Hicks brought Wister a gift of plants which included 61 accessions representing 3,143 individual plants. these included seven seedlings of the sweetbay magnolia (Magnolia viginiana), a native species which was then known as Magnolia glauca. of these original seven, only one survived. It was SCott ARBoREtum The original type specimen of Magnolia virginiana var. australis `Henry Hicks' still thrives at the Scott Arboretum (above). This cultivar bears fragrant, creamy white flowers and cold-hardy evergreen foliage (right). 4 Arnoldia 68\/1 I Early History of the Scott Arboretum n 1929, John Caspar Wister was appointed the first director of the Arthur Hoyt Scott Horticultural Foundation (now the Scott Arboretum). Wister graduated in 1909 with a degree from the School of Landscape Architecture at Harvard university, and supplemented this education with courses taken at the New Jersey Agricultural College. After graduation, Wister worked in landscape architecture offices in both Philadelphia and New York. From his youth, John Wister was an avid plant collector. As a small boy he had exposure to estate gardening at different Wister properties located in and around Germantown, Philadelphia. At age 14 he grew 40 cultivars of chrysanthemums. After Wister started his professional career his interest in a myriad of plant groups and genera began to grow. throughout his lifetime he was an avid collector of both herbaceous and tree peonies. Wister admired a photograph in a garden catalog that showed the peony collection of Arthur Hoyt Scott (for whom the Arboretum is named) and Edith Wilder Scott and in 1913 he met the Scotts at their home in oak Lane, Philadelphia. on July 10, 1917, at the age of 30, Wister enlisted as a private in World War I. Wister was sent to France. on his leave time during the war Wister toured the gardens of Europe. While in France he collected several cultivars of tree peonies and sent the plants back to mr. and mrs. Scott. Wister was honorably discharged in 1919. Arthur Hoyt Scott was a graduate of the class of 1895 from Swarthmore College. His father, E. Irvin Scott, founded Scott Paper Company which was located in Chester, just south of Swarthmore, Pennsylvania. Like Wister, Scott developed a passion for ornamental horticulture as a young man. In 1920 he became president of the Scott Paper Company, but his spare time was primarily occupied by his love of plants. Scott served as an officer of the American Peony Society and the American Iris Society. As early as 1915 Scott was sending gifts of plants to his alma mater, Swarthmore College. His first gift was 100 lilacs of many different varieties. In 1919 the Scotts moved from Philadelphia to a 100-acre farm in Rose Valley near Swarthmore. As Wister later wrote \"Here for the first time he had ample room. He at once began to plant great collections of flowering trees and shrubs like Japanese cherries, crabapples, dogwoods, lilacs, mockoranges and azaleas.\" When Arthur Hoyt Scott wanted to study peonies he had to travel to Cornell university and when he wanted to see lilacs he had to go to Highland Park in Rochester, New York. Scott dreamed of having an arboretum at Swarthmore College where local gardeners could go and see attractive displays of his favorite plants. Scott had the support of Samuel Palmer, the head of the Botany Department, and Swarthmore College. Palmer, in turn, contacted Robert Pyle who had graduated from Swarthmore in 1897 and was serving on Swarthmore's board of managers. Pyle was head of the Conard-Pyle Company, one of the country's largest purveyors of mail-order roses. Magnolias at the Scott Arboretum 5 John C. Wister (second from right) at the dedication of the Scott Arboretum's rose garden in 1958. Wister was director of the Scott Arboretum from 1929 to 1969. Arthur Hoyt Scott died in 1927, at the age of 51. two years later Edith Wilder Scott and Arthur Hoyt Scott's sister, margaret moon, and her husband, owen moon, approached Swarthmore's president with the idea of starting a campus arboretum. they recommended that John Wister become its first director, and so indeed he did. the early 1930s were the heydays of the Scott Horticultural Foundation. With Wister at the helm, the plant collections grew very quickly. Huge collections of Paeonia, Iris, Rhododendron, Syringa, Philadelphus, Prunus, Malus, Cotoneaster, Chrysanthemum, Narcissus and Magnolia were being accessioned and planted. In 1931 the Foundation accessioned 783 plants; in 1932 there were 1162 accessions, and in 1933, 1110 accessions. to put this in perspective the Scott Arboretum currently accessions about 300 plants per year. ARCHIVES oF tHE SCott ARBoREtum oF SWARtHmoRE CoLLEGE 6 Arnoldia 68\/1 NANCY RoSE planted in a poorly drained section of the magnolia Collection, and over the years this sweetbay magnolia thrived (unlike most magnolias, this species performs well in wet soils). It was observed that while most specimens of Magnolia viriginiana in the Swarthmore area are deciduous, this particular specimen was reliably evergreen. In 1967 this clone was officially registered and named Magnolia virginiana var. australis `Henry Hicks'. the original type specimen remains in great shape today in the old magnolia Collection. A Stream of Magnolias In addition to Magnolia virginiana, several accessions of oyama magnolia (Magnolia sieboldii, previously M. parviflora), a shrubby Asian magnolia noted for its white flowers with striking crimson stamens, were added to the collection from several different sources. other early additions included the star magnolia (Magnolia stellata), anise magnolia (Magnolia salicifolia), umbrella magnolia (Magnolia tripetala), Kobus magnolia (Magnolia kobus), southern magnolia (Magnolia grandiflora), cucumbertree magnolia (Magnolia acuminata), and the saucer magnolia (M. x soulangiana, syn. Magnolia x soulangeana). Magnolia x soulangiana resulted from a cross between Magnolia denudata and Magnolia liliiflora in 1820 by Etienne Soulange-Bodin, who was the first director of the Royal Institute of Horticulture near Paris. For many gardeners across the united States, saucer magnolia is the quintessential magnolia species. this large shrub to medium-sized tree produces masses of large, showy flowers that emerge before the foliage. the flowers, which are often fragrant, appear in white and shades of pink and purple. In the early 1930s the Scott Arboretum received two different batches of Magnolia x soulangiana cultivars. In 1933, Arthur D. Slavin at Highland Park in Rochester, New York, sent `Alexandrina', which has deep red-purple flowers and was introduced in Paris in 1831; `Amabilis', an 1865 French introduction with white flowers; `Alba', which is another white-flowered clone that was grown and named by Louis van Houtte of Belgium; `Andre Leroy', which has dark pink to purple flowers and is a French The slightly nodding flowers of Magnolia sieboldii bloom in late spring or early summer. NANCY RoSE Early-spring-flowering Magnolia salicifolia has fragrant, 6-tepaled white flowers and a pyramidal growth habit. Magnolias at the Scott Arboretum 7 SCott ARBoREtum introduction from 1892; `Brozzoni', which bears white flowers with pink veins and was named in honor of Camillo Brozzoni in Brescia, Italy in 1873; `Lennei', which has tepals that are magenta on the outside and white on the inside; `Norbertii', a lateblooming cultivar with red-purple flowers; and `Verbanica', which has deep pink flowers and was named by Andre Leroy in France in 1873. In 1936, scions of all these clones were sent to Verkades Nursery in Wayne, New Jersey. the magnolias were propagated there, and duplicate plants were then sent back to the Scott Arboretum. today, many of these original cultivars from Highland Park are found in our collections. Noted magnolia expert Philippe de Spoelberch from Arboretum Wespelaar, Haacht-Wespelaar, Belgium, commented that the Scott Arboretum's collection of Magnolia x soulangiana cultivars is important because they most likely represent clones which are true to name. De Spoelberch said that many of the original cultivars from France are much confused in the nursery industry and that many cultivar names have been mistakenly attributed to the wrong cultivar. Magnolia x soulangiana `Alexandrina' is noted for its dramatic redpurple flowers. A David Leach hybrid of M. acuminata x M. denudata, `Ivory Chalice' bears large, pale yellow to cream colored flowers. Southern Belles and Little Girls In 1933, the Scott Arboretum received its first plant of the southern magnolia (Magnolia grandiflora) as a gift from Edith Wilder Scott. this large magnolia, native to the southeastern united States, is prized for its leathery evergreen foliage and large, fragrant, creamy white flowers. Several cultivars of this species were soon added to the collection; in 1939, `Exoniensis' was received from Princeton Nursery, and in 1940 `Lanceolata' arrived from Hillier and Sons in Winchester, England. Both of these clonal names are synonymous with `Exmouth', which is a fastigiate cultivar. It was not until 28 years later, in 1968, that any additional selections of the southern magnolia were added to the Arboretum's collections. `Edith Bogue' was a selection that was made in 1961 for its ability to withstand very cold temperatures with minimal SCott ARBoREtum 8 Arnoldia 68\/1 NANCY RoSE SCott ARBoREtum Three of the \"Little Girl\" magnolia hybrids, (clockwise from above) `Betty', `Ann', and `Judy'. This group of magnolias was bred at the United States National Arboretum and named for the wives, daughters, and secretaries of the breeders. leaf burn. our plant came from Kingsville Nursery in Kingsville, maryland. today, there are several specimens of `Edith Bogue' growing on the campus of Swarthmore College, as well as 7 other M. grandiflora cultivars including both `D. D. Blanchard' and `Pocono' which also have been selected for greater cold hardiness. In 1968 the Scott Arboretum also received an important collection of magnolias from the united States National Arboretum. Commonly referred to as the Eight Little Girls, these magnolias were the result of hybridizing work conducted at the uSNA by research geneticist Dr. Francis deVos and horticulturist William Kosar. In 1955, deVos began breeding working using Magnolia liliiflora `Nigra' and Magnolia stellata `Rosea'. `Nigra' was used for its hardiness and late blooming, while `Rosea' was used for its fragrance, prolific flowering, and mildew resistance. the results of this program resulted in the introduction of cultivars `Ann', `Judy', `Randy', and `Ricki'. In 1956, Kosar hybridized Magnolia stellata `Rosea' and `Waterlily' with Magnolia liliiflora `Nigra' and `Reflorescens', which resulted in the introduction of cultivars `Betty', `Jane', `Pinkie', and `Susan'. today at NANCY RoSE Magnolias at the Scott Arboretum 9 SCott ARBoREtum A specimen of the rare Florida native Magnolia macrophylla subsp. ashei growing at the Scott Arboretum. the Scott Arboretum `Ann', `Betty', and `Susan' remain as beautiful mature specimens, while the others that we lost have been replaced with younger specimens. the \"Little Girl\" hybrids remain a group of magnolias that we continue to promote as relatively small (about 12 to 20 feet [3.5 to 6 meters] tall) magnolias for the home garden. In addition to Magnolia virginiana and Magnolia grandiflora, the Scott Arboretum added several other magnolia species native to the united States. We received the umbrella magnolia (Magnolia tripetala) from the Hicks Nursery in 1932 and Magnolia fraseri came from Arthur D. Slavin at Highland Park Nursery in 1933. Magnolia macrophylla, which is closely related to Magnolia fraseri, was acquired from Andorra Nursery near Philadelphia in 1939. the Scott Arboretum's first plant of Magnolia pyramidata (which is sometimes listed as Magnolia fraseri subsp. pyramidata) came to us via the Henry Foundation for Botanical Research in Gladwyne, Pennsylvania in 1971. this species is native to the coastal plains of Alabama, Georgia, Florida, Louisiana, mississippi, South Carolina, and texas, while Magnolia fraseri is only found in the mountains. It wasn't until 1991 that we added the last of the North American native magnolias, a single plant of Magnolia macrophylla subsp. ashei. Ashe's magnolia is very rare in the wild and only occurs in a small portion of the Florida panhandle where it is found from Leon to Wakulla counties and westward to Santa Rosa county. In the Red List of Magnoliaceae, which documents globally threatened plants within the magnolia family, Magnolia macrophylla subsp. ashei is given the conservation status of \"vulnerable\", which means it is considered to be facing a high risk of extinction in the wild. 10 Arnoldia 68\/1 CouRtESY oF PAt mCCRACKEN `Gold Crown', an August Kehr hybrid, bears large, light to medium yellow flowers. Magnolia zenii is a critically endangered species in its native range in China. Recent Additions, Future Plans the 1990s saw dozens of new cultivars enter the Scott Arboretum's collections from many magnolia purveyors such as Arbor Village Nursery, Gossler Farms, and Fairweather Gardens. In 1998, through Pat mcCracken and mcCracken Nursery, we received a number of cultivars introduced by noted magnolia hybridizer Dr. August Kehr. After retiring from the uSDA, Kehr started a robust magnolia breeding program in Hendersonville, North Carolina that resulted in many outstanding cultivars of magnolias. Some of the Kehr cultivars included in our magnolia collection are `Serenade', `Pink Perfection', and a number of the much-desired yellow-flowered hybrids including `Gold Crown', `Golden Endeavor', `Hot Flash', `Solar Flair', and `Sunburst'. to create the yellow magnolias Kehr made complex crosses using M. acuminata, M. denudata, M. x brooklynensis, M. `Elizabeth', M. `Woodsman' and M. `Gold Star'. From 2000 to 2010 the Scott Arboretum continued to add dozens of new magnolia taxa to our collection. many new cultivars of Magnolia grandiflora and Magnolia virginiana were added. Several other yellow-flowered magnolias such as `Yellow Joy', `Limelight' and `Golden Rain' were added. In addition, many species magnolias from a variety of sources were accessioned, including Magnolia x wiesneri, a hybrid between M. sieboldii and M. obovata; Magnolia zenii which is critically endangered in China where only one population, comprised of 18 NANCY RoSE Magnolias at the Scott Arboretum 11 SCott ARBoREtum Magnolia denudata `Swarthmore Sentinel' was selected and named for its distinctly upright habit. 12 Arnoldia 68\/1 SCott ARBoREtum individual trees, exists; and Magnolia wilsonii, which is endangered and only exists in scattered populations in Sichuan, northern Yunnan, and Guizhou, China. two other additions--Magnolia lotungensis from China and M. tamaulipana from northeastern mexico--may prove to be borderline hardy in Swarthmore (uSDA zone 6, average annual minimum temperature -10F to 0F [-23.3C to -17.8C]). In 2009 the Arboretum introduced a new selection of the Yulan magnolia, Magnolia denudata `Swarthmore Sentinel'. the Arboretum originally received a seedling from J. C. Raulston at North Carolina State university, who had received seeds from the Beijing Botanic Garden. From a seedling in 1993, the tree is over 30 feet tall today. on several occasions visiting magnolia experts commented on how upright our particular clone was. therefore, we decided to name this selection `Swarthmore Sentinel' for its fastigiate habit. over the last 81 years we have accessioned 502 magnolias at the Scott Arboretum. today the collection holds 165 different taxa. the Scott Arboretum's collection is recognized as a national magnolia collection through the American Public Garden Association's North American Plant Collections Consortium (NAPCC). According to the APGA \"the North American Plant Collections Consortium is a network of botanical gardens and arboreta working to coordinate a continent-wide approach to plant germplasm preservation, and to promote high standards of plant collections management.\" the Scott Arboretum will be working with approximately 20 other institutions across North America, including San Francisco Botanical Garden, Quarryhill Botanical Garden, university of British Columbia Botanical Garden, the Bartlett Arboretum, and Atlanta Botanical Garden to create a consortium of institutions to oversee the preservation and conservation of magnoliaceae germplasm. this group will also be part of the NAPCC and administered through the APGA. once formed, this magnolia Curatorial Group will partner with the magnolia Society International to target both wild species and cultivar groups which need to be preserved in botanic gardens and arboreta. the Scott Arboretum Magnolia `Charles Coates' is an unusual hybrid between M. sieboldii and M. tripetala. will also continue to grow its own collections. We currently have 72 magnolia taxa growing in a nursery, and once these reach specimen size they will be transplanted to garden sites throughout the arboretum. In 2015 the Scott Arboretum plans to host the international meeting of the magnolia Society International. Bibliography Callaway, D.J. 1994. The World of Magnolias. timber Press, Portland, oregon. Gardiner, J. 2000. Magnolias: A Gardener's Guide. timber Press, Portland, oregon. Liu, Y.H. 2004. Magnolias of China. Hong-Kong, Beijing Science & technology Press. treseder, N.G. 1978. Magnolias. Faber and Faber, Limited, London and Boston. Wister, J.C. Swarthmore Plant Notes 19301954, Volume 1, Part 1. pp. 8088. Yagoda, B. 2003. The Scott Arboretum of Swarthmore College--The First 75 Years. the Donning Company Press. Andrew Bunting is Curator of the Scott Arboretum of Swarthmore College in Swarthmore, Pennsylvania. "},{"has_event_date":0,"type":"arnoldia","title":"Excerpts from Wild Urban Plants of the Northeast: A Field Guide","article_sequence":2,"start_page":13,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25482","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060a76b.jpg","volume":68,"issue_number":1,"year":2010,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Excerpts from Wild Urban Plants of the Northeast: A Field Guide Peter Del Tredici E Editor's NotE: ver wonder what kind of tree that is, the one growing from a crack in the asphalt parking lot at work? Or what that tangled vine engulfing the slope by the subway station might be? Wild Urban Plants of the Northeast: A Field Guide, written by long-time Arnold Arboretum researcher Peter Del Tredici, may have your answer. Del Tredici's goal with this book is \"to help the general reader identify plants growing spontaneously in the urban environment and to develop an appreciation of the role they play in making our cities more livable.\" Many of the 222 plants featured in the book could be called weeds, and some are notoriously invasive. The author eschews these labels, however, pointing out that in many urban\/suburban areas the environment has been so radically altered (think non-native fill soils, soil compaction and contamination, impermeable pavement, and pollution) that the presence of any plants has benefits. This handy guide is organized by plant families and includes both woody and herbaceous plants. Numerous color photographs and extensive information is provided for each species, including place of origin, descriptions of vegetative, flower, and fruit characteristics, and habitat preference. Some fascinating details emerge from the \"Cultural Significance\" subsections--for example: \"During World War II, the silky seed hairs [of common milkweed, Asclepias syriaca] were used as a substitute for kapok to fill \"Mae West\" life vests. Between 1943 and 1945, a million such flotation devices were filled with the floss from some 24 million pounds (11 million kilograms) of milkweed pods.\" Following are half a dozen plant species featured in the book. Reprinted from: Peter Del Tredici, Wild Urban Plants of the Northeast: A Field Guide. Copyright 2010 by Cornell University. Used by permission of the publisher, Cornell University Press. 374 pages. ISBN 978-0-8014-7458-3. 14 Arnoldia 68\/1 Wild Urban Plants of the Northeast 15 16 Arnoldia 68\/1 Wild Urban Plants of the Northeast 17 18 Arnoldia 68\/1 Wild Urban Plants of the Northeast 19 20 Arnoldia 68\/1 Wild Urban Plants of the Northeast 21 22 Arnoldia 68\/1 Wild Urban Plants of the Northeast 23 24 Arnoldia 68\/1 Wild Urban Plants of the Northeast 25 "},{"has_event_date":0,"type":"arnoldia","title":"Conserving the Dawn Redwood: The Ex Situ Collection at the Dawes Arboretum","article_sequence":3,"start_page":26,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25481","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060a726.jpg","volume":68,"issue_number":1,"year":2010,"series":null,"season":null,"authors":"Payton, Greg","article_content":"Conserving the Dawn Redwood: The Ex Situ Collection at the Dawes Arboretum Greg Payton S NANCy ROSE ince 1990, the Dawes Arboretum in Newark, Ohio, has undertaken a large scale ex situ conservation project with Metasequoia glyptostroboides, the dawn redwood. Ex situ conservation is defined as the conservation of genes or genotypes outside their environment of natural occurrence (China, in the case of dawn redwood). There are challenges and limits to ex situ conservation, but for some threatened or endangered plants and animals it is an essential component in efforts to keep the species from extinction. For a long-term conservation project to be successful and sustainable, a large sampling of genetic material is desirable to maintain the existing and potential variation within a particular species. Many attempts at rescue efforts are done on a limited basis, and they hold relatively small numbers of specimens due to insufficient space and budgetary limitations. Ideally, ex situ collections should have the capacity to grow the requisite number of individuals essential for preserving the base Dawn redwoods develop distinctive buttressed trunks with age. Dawn Redwood at the Dawes Arboretum 27 GREG PAyTON GREG PAyTON A specimen with good form and foliage qualities (accession D1993-0249.004). Variations in foliage of trees in the Dawes plantation. All branchlets photographed on October 13, 2009. gene reserve with a goal of capturing as large of a part of the genetic diversity within the species as possible. Some species require relatively few individuals to capture that genetic range, while others require much larger population sizes. Studies of the genetic variation within dawn redwood have been and still are being conducted. Early results indicate that there is a fairly low genetic diversity, although there is some differentiation within the native populations throughout the overall range of the species. Ex situ conservation does have its limits, and ideally it should complement in situ protection in the natural environment. Preserving a native, wild population is the best option, and this should be the primary focus of any conservation program. One of the particular problems with ex situ conservation lies in the inevitable environmental differences between the site of origin and the site of the ex situ collection. If plants in the ex situ site are allowed to sexually reproduce, environmental conditions in this new setting favor the selection and survival of the progeny best adapted to that site. Progeny that survive in the ex situ location may have different traits than progeny which would have survived in the original site. While this may have advantages from a horticultural standpoint (e.g. selection of plants with greater cold hardiness or better drought tolerance), it is a disadvantage for most conservation goals. Preserving the genetic diversity of a species ex situ may be best accomplished by maintaining clonal populations. However, seed banking of species with orthodox seeds (seeds that survive drying or freezing) can also be important in securing a species for the future, and there is the advantage that seeds can be stored in a much smaller space than living plants. A combination of both seed banking and living plants offers the most opportunities for conservation research. 28 Arnoldia 68\/1 Meet Metasequoia glyptostroboides Dawn redwood (shui-shan in Chinese, meaning \"water-fir\") is a deciduous conifer similar to bald cypress (Taxodium distichum). The soft, distichous needles of dawn redwood are arranged oppositely, easily distinguishing it from bald cypress with its alternate needle arrangement. When dawn redwood--once thought to be extinct--was discovered still growing in southcentral China in a mild and wet climate, it was not believed that it would survive in the United States north of Georgia. The provenance testing done since Metasequoia seeds arrived in the United States in 1948 shows that it can survive in USDA Hardiness Zones 5 to 8 (average annual minimum temperature -20 to 20F [-28.8 to -6.7C]) in areas with sufficient rainfall (or with supplemental watering). In its native Sichuan, China, the average rainfall is around 40 inches (100 centimeters) per year but dawn redwood has survived in parts of the United States with lesser amounts of rainfall. The typical form is a large tree, up to 150 feet (45 meters) tall in the wild, pyramidal in youth, becoming more open-crowned with great age. The trunks on older specimens become strongly buttressed. It is fast growing when moisture is available and can add over 3 feet (1 meter) of growth per year. It is heliophilic (requiring full sun), which has limited its use as a commercial timber tree since it does not grow well in competition. Many millions of dawn redwoods have now been planted throughout China, but the condition of the native population has remained stagnant. The 2009 IUCN Red List of Threatened Species gives dawn redwood a status of critically endangered, saying that the few remaining trees have been protected but that the habitat has not been, and there are poor prospects for natural regeneration. The valleys the tree prefers have been denuded of vegetation and mature trees are often limbed up--all the way to the top--for firewood. Seedling reproduction is unlikely in this altered environment. In the past, natural seeding was also hampered because the seeds were collected and sold by farmers for various uses such as timber plantations. This practice has become less common in recent years, since An example of a plant that exists only ex situ is Franklinia alatamaha, Franklin tree. It is believed to have been extirpated from its native range (Georgia, in the southeastern United States) by the early nineteenth century. Fortunately, botanists John and William Bartram found and later collected and propagated Franklin tree in the late eighteenth century, and the species still survives in cultivation today. It blooms from late summer into autumn, and flowering often overlaps with fall foliage color. other Chinese conifers have provided lumber of greater quality. In addition, propagation from cuttings has proven to be advantageous for producing new plants. Recent surveys indicate that 5,396 native trees (of all ages) still remain in the native range in China. The majority of trees (5,363) grow in western Hubei, while 28 grow in eastern Chongqing. Only 5 trees remain in Hunan. NANCy ROSE Dawn Redwood at the Dawes Arboretum 29 A Case of Depression and the \"Single Tree\" Theory In 1983, Dr. John Kuser, a forestry professor at Rutgers University, surmised that cultivated Metasequoia in the United States were suffering from inbreeding depression. He said, \"Apparently, variation in the amount of genetic load carried by different trees causes some to be incapable of producing fertile selfpollinated seeds but allows others to produce a few viable seeds and occasional trees to self quite well.\" He noted that Metasequoia pollen is wingless and \"tends to clump together.\" The best seed germination was found to occur on trees that had been located advantageously for cross-pollination. At the time, the popular belief was that the poor germination of seedlings was the result of trees in the United States having all originated from the single \"type\" tree in the village of Maudao, China. However, allozyme variation work done in 1995 showed that the 1947 seeds were not likely to have come from a single isolated tree. Furthermore, a copy of a previously unpublished paper by W. C. Cheng dated March 25, 1948 revealed, as stated above, that Hwa had found more than 1000 Metasequoia and about 100 \"big ones.\" Apparently seeds from many MAP by GREG PAyTON Map of native dawn redwood distribution and seedlot collection sites. 30 Arnoldia 68\/1 GREG PAyTON trees had been collected and disseminated. Poor seed set seems to stem from the fact that most seed production outside of China is the result of selfing (due to isolation of specimens). The genetic variation of dawn redwood in China was believed to be much greater than that in the United States, and in 1990 a cooperative research project on Metasequoia began between Dr. W. J. Libby at the University of California, berkeley, Professor Minghe Li at Huazhong Agricultural University in Hubei, China, and Dr. Kuser. A number of organizations contributed to fund the project, and it was at this point that the Dawes Arboretum became involved in provenance testing of Metasequoia. Professor Li collected Metasequoia seeds from several locations in its native range in October 1990. In April 1991, 53 packets of seeds were PHOTO by bURNEy HUFF (DAWES ARCHIvES) A bronze-foliaged specimen in the plantation (accession D1993-0237.005). received at Rutgers University from Professor Li, 52 from trees that still had seed cones, and one packet of mixed seeds. These seed lots were germinated, and only four of the collections produced no seedlings. The remaining 48 \"families\" were grown on, and complete collections were planned for both Rutgers and Dawes. The remaining seedlings were distributed to nearly 20 cooperating institutions and individuals in the United States and United Kingdom. (The Arnold Arboretum received 125 of these seedlings.) In 1993 the Dawes Arboretum received two shipments of the dawn redwood seedlings from Rutgers. A total of 344 trees were planted in the Dawes plantation. because of the large size (8 acres [3.2 hectares]) of the Dawes site we were able to plant the trees 25 feet (7.6 meters) apart so no subsequent thinning was necessary. A dawn redwood specimen from the original 1949 seed accession at the Dawes Arboretum. The photograph is from the early 1990s when the tree was nearly 80 feet (24 meters) tall; a lightning strike later took out the top of the tree. Current Status of the Dawes Collection The Dawes plantation of seedlings from Professor Li and Rutgers currently consists of 320 trees, which makes it one of the largest living ex situ conservation collections of documented wild-origin dawn redwood trees outside of China. Through 2009, 24 trees have been lost Dawn Redwood at the Dawes Arboretum 31 from this plantation, and one seed lot family has been lost completely from both the Dawes and Rutgers plantations. In 2009, Dawes began contacting other institutions to see what living accessions they had from the original 52 seed lots; 29 new accessions (in the form of vegetative cuttings) representing trees from seed lots where Dawes had few representatives were obtained from these institutions. Since each of these trees was originally grown from seeds, every tree is genetically unique and therefore valuable for its individuality. These cuttings are currently doing well in propagation and will help to provide more genetic stock to add to the diversity of the plantation. The search for additional collections of this Li\/Rutgers project is ongoing. Any other modern or historical collection of wild material would be invaluable to add to the Dawes collection. One of the seed lots that had no germination was the only lot from Hunan, collected from three individual dawn redwoods there, so we are especially interested in acquiring germplasm from the few trees in Hunan. In addition to the plantation trees, Dawes has a few other accessions of wild-collected Metase- quoia: three accessions from the original 1947 seedlings, received in 1950 from Ralph Chaney who presumably got his seeds from Merrill; a grove of 44 trees propagated by cuttings in 1960 from the previous accession; and three individuals also propagated from the original accession. Into the Future In Metasequoia, female cones (macrosporangiate strobili) are typically produced when trees reach a height of 30 to 50 feet (9 to 15 meters ). Male cones (microsporangiate strobili) are not produced until trees are 60 to 83 feet (18 to 25 meters) in height. At this point, neither female nor male cones have been observed on the Dawes Arboretum plantation trees. As the grove continues to grow and seed production begins, the resultant progeny will represent the greatest level of genetic variation within dawn redwood outside of China. The origins of these plantation trees are from across the estimated 800 square kilometer (312 sq. mi.) native range in central China where full cross-pollination is very unlikely. Studies have shown that trees in the native populations show a lack of spatial genetic flow, indicating GREG PAyTON Wide spacing allows ample room for trees in the dawn redwood plantation. 32 Arnoldia 68\/1 DAvID bRANDENbURG The author with a witches'-broom on one of the Dawes plantation trees. In 2009, both the genetic and taxonomic (cultivar) collections of dawn redwoods at the Dawes Arboretum were granted full status as a North American Plant Collections Consortium (NAPCC) collection. This symbolizes the commitment of the staff and organization to fulfilling the duty of preserving this important collection. As a repository for North America, requests for propagation material are honored for research purposes. Of horticultural interest, there are well over two dozen cultivars of Metasequoia that add to the range of variation within the species. `Miss Grace' and `Bonsai' are dwarf selections, `Jack Frost' has a hint of variegation, and `Ogon' (syn. `Gold Rush') is a Japanese cultivar with bright yellow foliage that originated from irradiated seeds. Several cultivar selections could be made from the Dawes plantation trees, as there are some interesting habits and foliage types. Tree heights of the plantation trees are from scarcely 3 feet (1 meter) tall to over 33 feet (10 meters), and habits range from squat and round to tall and narrow with many forms in between. Foliage varies from large and coarse to small and fine, with colors in shades of green and bronze. A witches'-broom--which may yield dwarf forms--has even been found on one specimen. Dawn Redwood at the Dawes Arboretum 33 genetic isolation due to habitat fragmentation (Leng et al. 2007). As stated earlier, natural pollen dissemination is limited. Since these wide-ranging Chinese collections are located together in a single plantation at Dawes, broad genetic combinations could occur. The resultant mixed, open-pollinated seeds could prove useful for horticultural purposes as well as for selecting for resistance to any future insect or disease pressures. These seeds would have limited use for some conservation projects (since they are from mixed meta-populations), but there is potential for controlled crossing within the separate seed lot collections, which would give greater conservation value. The seeds produced here will be made available to seed banks, researchers, and growers. This collection holds many opportunities for future studies and research to be conducted without traveling to China. The sister population at Rutgers University is currently the subject of an amplified fragment length polymorphism (AFLP) analysis to assess the breadth of the genetic diversity of the collection. Since most of the genotypes at Rutgers are duplicates of dawn redwoods in the collection at Dawes, the data from the AFLP study will pertain to this collection as well. We hope that this successful ex situ collection at the Dawes Arboretum will aid in the conservation and further understanding of this ancient and impressive species. Bibliography Andrews, H.N. 1948. Metasequoia and the Living Fossils. Missouri Botanical Garden Bulletin 36(5): 7985. bartholomew, b., D.E. boufford, and S.A. Spongberg, 1983. Metasequoia glyptostroboides--Its Present Status in Central China. Journal of the Arnold Arboretum 64: 105128. Ecker, Eisenman, S.W. 2009. Pers. comm. Rutgers University, School of Environmental and biological Sciences, Department of Plant biology and Pathology. GSPC. 2002. Global Strategy for Plant Conservation. Montreal: Secretariat of the Convention on biological Diversity. Hendricks, D.R. 1995. Metasequoia Depression, Sex, and Other Useful Information. Landscape Plant News 6(2): 710. Hendricks, D. and P. Sondergaard. 1998. Metasequoia glyptostroboides--50 years out of China. Observations from the United States and Denmark. Dansk Dendrologisk Arsskrift 6: 624. Hsueh, C.-J. 1985. Reminiscences of Collecting the Type Specimens of Metasequoia glyptostroboides. Arnoldia 45(4): 1018. Hu, H.H. 1948. How Metasequoia, the \"living fossil\" was discovered in China. Journal of the New York Botanical Garden 49(585): 201207. IUCN. IUCN Red List of Threatened Species. Version 2009.1. Retrieved October 12, 2009, from www. iucnredlist.org Kuser, J.E., D.L.Sheely, and D.R. Hendricks. 1997. Genetic variation in Two ex situ Collections of the Rare Metasequoia glyptostroboides (Cupressaceae). Silvae Genetica 46(5): 258264. Kuser, J. 1983. Inbreeding Depression in Metasequoia. Journal of the Arnold Arboretum 64: 475481. Leng, Q. et.al. 2007. Database of Native Metasequoia glyptostroboides Trees in China based on New Census Surveys and Expeditions. Bulletin of the Peabody Museum of Natural History 48(2): 185233. LePage, b.A., C.J. Williams, and H. yang. 2005. The Geobiology and Ecology of Metasequoia. Springer. Li, M. 2009, November 1. Pers. comm. Li, X.-D., H.-W. Huang, and J.-Q. Li. 2003. Genetic diversity of the relict plant Metasequoia glyptostroboides. Biodiversity Science 11: 100108. Ma, J. 2003. On the unsolved mystery of Metasequioa. Acta Botanica Yunnanica (25)2: 155172. Ma, J. 2003. The Chronology of the \"Living Fossil\" Metasequoia glyptostroboides (Taxodiaceae): A Review (19432003). Harvard Papers in Botany 8(1): 918. Ma, J. 2002. The History of the Discovery and Initial Seed Dissemination of the Metasequoia glyptostroboides, A \"Living Fossil\". Aliso 21(2): 6575. Ma, J. and G. Shao. 2003. Rediscovery of the \"first collection\" of the `Living Fossil', Metasequoia glyptostroboides. Taxon 52(3): 585588. Merrill, E.D. 19981999. Another Living Fossil Comes to the Arnold Arboretum. Arnoldia 5859(4-1): 1719. Sand, S. 1992. The Dawn Redwood. American Horticulturist 71(10): 4044. Wyman, D. 1968. Metasequoia After Twenty years in Cultivation. Arnoldia 28(1011): 113122. Greg Payton is the Plant Records Specialist at the Dawes Arboretum in Newark, Ohio. "},{"has_event_date":0,"type":"arnoldia","title":"Index to Arnoldia Volume 67","article_sequence":4,"start_page":34,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25483","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060ab6f.jpg","volume":68,"issue_number":1,"year":2010,"series":null,"season":null,"authors":null,"article_content":"Index to Arnoldia Volume 67 Items in boldface refer to illustrations Anteater 2: 30 Anthocyanins 3: 23 Anticancer plants 3: 23, 25 Antioxidant fruit 3: 1425 -- -- commercial potential of 3: 2325 Ants, leaf-cutter 2: 30 Appalachian Mts., Tennessee 3: 20 Apple, original 2: 20 -- fruiting genotypes 2: 20 -- quince and 1: 3 -- scab resistance 2: 10, 10, 20 Apple-pear, Asian 4: 28 Apomixis 3: 19, 21, 22, 2425 Arboriculture and plant hormones 4: 1519 Arborvitae, as beetle host 1: 35 Arisaema dilatatum, in China 2: 27, 28 Armenia, quince-growing in 1: 5, 5 Arnold Arboretum, Acer rubrum `Schlesingeri' at 2: 32, inside back cover -- -- aerial photographs of 1: 1927, 11; 1929, 13; 1936, 14; 1955, 13, 18; 1967, 12; 1968, 15; 2005, front cover, 11, 15; 2006, 18; 2007, 12; 2008, 17; 2009, 19; 2: 2008, 14 -- -- apple selection at 2: 20 -- -- autumn interest 2: 32, inside back cover; 4: 23 -- -- beetle research at 1: 3135, 32 -- -- Bentham and Hooker sequence at 2: 16 -- -- Bradley Rosaceous Collection 1: 14, 44; 2: 16, 20, 20; 4: 22, 24 -- -- Bussey Brook Meadow, in aerial photo 1: 14 -- -- Bussey Hill, in aerial photos 1: 11, 13, 14 -- -- Camellia trials 1: 27 -- -- cartography systems 1: 1219 -- -- Centre Street, in aerial photo 1: 14 -- -- China expeditions, 19071908, 1910 3: 213 -- -- cold-hardiness at 3: 36 -- -- conifer collection 3: 36; 4: 22 -- -- crabapple legacy 2: 1421, back cover -- -- Crataegus at 2: 16 A Abies spp., and exotic beetles 1: 33, 35 -- homolepis, lightning-damaged 4: 22, 22 Abscisic acid 4: 15, 1819 -- -- photosynthesis and 4: 19 Acai juice 3: 23 Acer spp., and exotic beetles 1: 35 -- davidii, in China 2: 22, 26 -- -- bark 2: inside front cover -- rubrum `Schlesingeri' 2: 32, inside back cover -- -- -- propagation and redistribution of 2: 32 -- saccharum 3: 31 -- sutchuenense, in China 2: 27 Ackerman, Dr. William 1: 24, 28 Acorns, features of 4: 25, 35, 10, 11 Adenorachis 3: 21 Aerial photography and mapping 1: 1019, 1115, 1719 Aesculus spp., and exotic beetles 1: 34, 35 Afghanistan, pine from 3: 36, inside back cover Africa, pest beetles from 1: 33 Agrilus planipennis 1: 34, 34 Agroforestry 3: 2627 Aiello, Anthony S., \"Seeking ColdHardy Camellias\" 1: 2030 Ailuropoda melanoleuca, discovery of 2: 23 Akebia trifoliata, in China 2: 26 Alders, as beetle host 1: 35 Alexander, John H., III -- -- -- -- photographs by 1: inside front\/back covers; 2: 18 Allium tricoccum 3: 30 Alnus spp., and exotic beetles 1: 35 Alpha-pinene 1: 32 Alpine plants, in China 3: 213, 4, 6, 1011 Ambrosiella fungi 1: 35 American ginseng 3: 2830, 2930, 35 Amplified fragment length polymorphism (AFLP) 4: 7, 910 Animal and Plant Health Inspection Service (APHIS), and beetles 1: 3135 Anoplophora glabripennis 1: 34, 34 -- -- cultivar evaluation 2: 18 -- -- Dana Greenhouses, in aerial photos 1: 14 -- -- early accessions 1: 44; 2: 16, 1920 -- -- Faxon Pond 2: 32 -- -- Forest Hills Gate 2: 16 -- -- Forsythia hybrids at 2: 18 -- -- Himalayan pine at 3: 36, inside back cover -- -- Hunnewell building, in aerial photos 1: 15, 15 -- -- Hydrangea paniculata `Praecox' at 1: inside covers, 44 -- -- introductions 1: 44; 2: 6, 1821 -- -- Japanese and Korean plants at 1: 27, 44; 2: 16 -- -- Leventritt Shrub and Vine Garden, aerial photo of 1: front cover -- -- Living Collections survey 1: 15, 17 -- -- Master Plan 1: 17 -- -- Meadow Road 2: 32 -- -- Malus collection 2: 4, 1421, 14, 1621, back cover -- -- Metasequoia glyptostroboides at 4: 23 -- -- model 1: 1819 -- -- Nikko fir, loss of 4: 22, 22 -- -- Peters Hill 2: 14, 16, 16, 18, 19, 19, 20 -- -- -- -- in aerial photos 1: 1112, 14 -- -- Pinus wallichiana at 3: 36, inside back cover -- -- plant distribution benefits 2: 20 -- -- Prunus at 1: 13; 2: 18; 4: 24 -- -- Pyrus 2: 16 -- -- -- pyrifolia at 4: inside covers, 28 -- -- Rosaceae blights at 4: 22 -- -- sand pear at 4: 28, inside back cover -- -- Visiting Committee, 1955 1: 15 -- -- \"Weather Station Data--2009\" 4: 2024 -- -- Weld Hill in aerial photos 1: 14, 18, 19, 19 -- -- winter temperatures 3: 36 Arnoldia, Index to Volume 66 1: 3643 -- and Donald Wyman 2: 19 Index 35 Arnot Teaching and Research Forest 3: 32, 32 Aronia 3: front\/back covers, 1425, 1418, 22, 24 -- arbutifolia 3: 1415, 1519, 2122, 24 -- -- flowers 3: 15 -- -- foliage 3: 15 -- -- fruit 3: 14 -- fruit chemistry 3: 14, 21, 2325 -- -- crop potential 3: 1925 -- genetics 3: 19, 21, 22, 2425 -- habitat and range 3: 1819, 1820, 21 -- hybrids 3: 21, 25 -- juice products 3: 2225, 23 -- `Likernaya' 3: 25 -- melanocarpa 3: 1519, 16, 17, 2125, 22, 24 -- -- foliage 3: 17, 22 -- -- fruit 3: front cover, 16, 24 -- -- growth habit 3: 21, 22 -- -- `Nero' 3: 24, 25 -- -- x Sorbus aucuparia 3: 25 -- -- `Viking' 3: 24, 25 -- mitschurini 3: 25 -- ploidy and apomixis in 3: 19, 21, 22, 2425 -- prunifolia 3: 15, 1819, 21, 22 -- -- x arbutifolia 3: 21 -- -- x melanocarpa 3: 21 -- -- x prunifolia 3: 21 -- taxonomy 3: 21 -- -- and Photinia 3: 21 \"Aronia: Native shrubs With Untapped Potential,\" Mark Brand 3: 1425, 1420, 2224 Ash, as beetle host 1: 34 -- borer, emerald 1: 34 Asia, plants from 1: 2030, 44; 2: 5, 2228; 3: 213, 36; 4: 28 Asian long-horned beetle (ALB) 1: 34, 34; 2: 29 -- medicine, traditional 3: 2930 Asiatica Nursery [PA] 1: 2021 Asimina triloba, fruit of 3: 28, 28, 30 Astilbes, shade-grown 3: 33 Atomic testing 2: 31 Autumn color 2: 32; 4: 23, 28 \"Autumn's Harbinger: Acer Rubrum `Schlesingeri',\" Michael S. Dosmann 2: 32, inside back cover Auxin pathway 4: 1519 -- exogenous 4: 18 B Bachtell, Kris, photo by 2: inside front cover Bacterial diseases 2: 10; 4: 22 Bamboo, in panda habitat 2: 26 Baoxing, plant exploring in 2: 2228 Bark beetles, in port of Boston 1: 3132 Basset, Cedric,\"In the Footsteps of Father David\" 2: 2228, 2228 Bayesian approach 4: 11 Beech 3: 31 Beeches, as beetle host 1: 35 Beetle, ambrosia 1: 32 -- Asian long-horned (ALB) 1: 34, 34; 2: 29 -- emerald ash borer (EAB) 1: 34, 34 -- European spruce bark 1: 35 -- red-haired pine bark 1: 33, 33 -- six-toothed bark 1: 33, 33 Beetles, damaging 1: 3135, 334 -- -- emergence and phenology 1: 32, 35 -- -- fungal vectors of 1: 33, 35 -- -- links to information 1: 34 -- -- new surveys and trapping methods 1: 3235 -- -- observation of 1: 34 Bene, John 3: 27 Bentham, George 4: 26 Bentham and Hooker sequence 2: 16 Berberidaceae 2: 26 Beresowski (the botanist) 2: 28 Berks, Robert 4: 27 Berry crops 3: 1425, 28, 30 \"`Best' Crabapples (Malus spp.)\" 2: chart 9 Betula spp., and exotic beetles 1: 35 \"Between Earth and Sky: Our Intimate Connections to Trees,\" Nalini M. Nadkarni, [excerpt] 2: 2931 Bible, quince in 1: 3 Binomial nomenclature 4: 26 Biodiversity 2: 2223, 24, 28; 3: 6, 1113, 26, 27, 28 Biology and taxonomy 4: 2527 Birch spp. 3: 36 Birches, as beetle host 1: 34, 35 Birds 2: 6, 10; 3: 14, 16 \"Bird's-eye Views: Aerial Photographs of the Arnold Arboretum,\" Sheila Connor 1: 1019, 1019 Black, James W., aerial photography of 1: 10, 10 Blackberries 3: 28 Blights and 2009 weather 4: 20, 22 Blooming, premature 4: 24 Blue Ridge Community College 4: 19 Blue stain fungi 1: 33 Boston 133 Cities Urban Area mapping program 1: 17 Boston port 1: 31 -- -- invasive beetles and 1: 3132 Botryosphaeria obtusa 2: 10 Bourg, Ian C., Ph.D. 2: 28 Brand, Mark, \"Aronia: \"Native shrubs With Untapped Potential\" 3: 1425 -- -- photos by 3: back cover Bristol, Peter 1: 21 Brooklyn Botanic Garden 2: 6 Buckeyes, as beetle host 1: 34, 35 Burbank, Luther, and quinces 1: 2, 4, 7, 8 Burma, pine from 3: 36 Burnett, D. Graham 4: 27 Bussey Institute 2: 17 C Calanthe tricarinata, in China 2: 25, 25 California, quinces in 1: 2, 3 Callicarpa japonica 1: 24 \"Camellia Belt\" 1: 27 Camellia japonica 1: 2030, 2223, 2529 -- -- `Balustrade' 1: 29 -- -- `Bloomfield' 1: 29, 29 -- -- espaliered 1: 30 -- -- fruit and seed 1: 23 -- -- grazing and 1: 23 -- -- `Korean Fire' 1: 28 -- -- Korean selections 1: 2429, 27, 28, 29 -- -- `Longwood Centennial' 1: 28 -- -- `Longwood Valentine' 1: 28 -- -- `Meadowbrook' 1: 29 -- -- `Morris Mercury' 1: 29 -- -- winter performance 1: 20, 2430 Camellias, hardier 1: 2030 Campanulaceae 2: 26 36 Arnoldia 68\/1 Campbell, Nichole K., \"Searching for Exotic Beetles\" 1: 3135 Canada, ginseng cultivation in 3: 30 Cancer, plants against 3: 23, 25 Caprifoliaceae 2: 25 Cardamine, in China 2: 28 Cargo ships, and pests 1: 3132, 31 Carnegie Museum of Natural History, beetle collections 1: 32 Carpinus caroliniana 3: 31 -- fangiana, in China 2: 26, 27 Carya spp. 3: 30 -- x dunbarii 3: 32 -- graft unions 3: 32, 32 -- laciniosa x ovata 3: 32 -- ovata 3: 32 -- ovalis 3: 32 Caucasus region, quinces of 1: 4, 5, 7, 7 Cedars, as term 2: 23 Ceratocystis spp. 1: 33 -- polonica 1: 35 C-glucoside vitexin flavone 3: 21 Chaenomeles sinensis 1: 8 Chanticleer garden 1: 27 Chengdu Institute of Botany 2: 28 Cherries, as beetle host 1: 35 Cherry, flowering 4: 24 -- in secondary growth 3: 32 Chicago area, oaks near 4: 4, 7, 10 Chicago Botanic Garden 4: 27 China, expedition to, 1980 2: 19 -- Imperial 3: 11 -- travel between Tibet and 3: 213 -- plants of 1: 30; 2: inside front cover, 15, 16, 1920, 2228, 2223, 2528; 3: inside front cover, 213, 4, 6, 10, 11; 4: 28 Chinese medicine, traditional 3: 2930 Chokeberry, black 3: front cover, 1519, 16, 17, 2125, 22, 24 -- commercial potential of 3: 2225 -- comparison of spp. 3: 1418, chart 17, 19, 2123, back cover -- cultural needs 3: 19, 23 -- fruits 3: front cover, 1425, 14, 16, 24, back cover -- habitat and distribution 3: 1820, 1820 -- ornamental qualities 3: 1419, 1417, 22 -- populations 3: 21 -- propagation 3: 14, 22 -- purple 3: 1819, 21, 22 -- red 3: 1415, 1519, 21, 22, 24 -- winter interest 3: front cover Chagga people 3: 27 Chloroplast data, and oak hybrids 4: 3 Classification trends 4: 2527 Clematis, in China 2: 28 Climbing plants 2: 26, 27 Climate change 4: 13 Codonopsis tangshen, in China 2: 26 Cold-hardiness 3: 36 -- -- in camellias 1: 2030 Computerized records, and mapping 1: 17, 19 Conifers 3: 36; 4: 22 -- pests of 1: 3135 Connor, Jay 1: 19 -- -- photographs by 1: front cover Connor, Sheila, \"Bird's-eye Views: Aerial Photographs of the Arnold Arboretum\" 1: 1019 Container-growing 3: 33 Convallariaceae 2: 25 Corkscrew willow 3: 35 Cornell University 3: 32; 4: 25 Cornus controversa, in China 2: 25, 26 -- kousa, Korean 1: 30 -- sericea, in winter 3: 34 Corydalis anthriscifolia, in China 2: 26 -- davidii, in China 2: 26 Cotoneaster moupinensis, in China 2: 28 Crandall, C.S. 2: 20 \"Crabapple Cultivars Introduced by Arboretum\" 2: chart 21 Crabapple cultivars 2: 213, 1721 Crabapples, at Arboretum 2: 1421 -- bark interest 2: 1920 -- best of 2: chart 9 -- breeding and selection 2: 57, 1421 -- choosing 613 -- cultivation 2: 37 -- description 2: 25 -- diseases 2: 10, 20 -- fruit 2: 311, 4, 5, 8, 10, 11, 13, 17, 1920, 19 -- -- palatability of 2: 10 -- longevity 2: 5 -- plumleaf 2: 56 -- problems with 2: 23, 6, 7, 10, 13 -- seasonal interest 2: 5, 10, 13, 19 -- Siberian 2: 1920 -- siting 2: 3, 5, 67, 10, 13 -- weeping 2: 5, 56, 12, 13, 13 \"Crabapples...With No Apologies,\" Jeff Iles 2: 213, 28, chart 9, 1013 Crops, new 3: 1425, 2635 -- -- medicinal 3: 23, 25, 2930 -- -- ornamental 3: 3335 Cypripedium tibeticum 3: 12 Cryptomeria spp., and exotic beetles 1: 35 Cunninghamia lanceolata 2: 23, 23 Cuppressaceae 2: 23 Curtis, Ralph, 1922 photo by 2: 16 Cydomalus 1: 4 Cydonia oblonga 1: 29, 27, back cover -- \"A\" clone 1: 5 -- -- `Angers' 1: 5 -- -- botany of 1: 34, 8 -- \"C\" clone 1: 5 -- -- cultivation and uses 1: 27 -- -- germplasm resources 1: 37, 9 -- -- `Champion', 1909 illustration 1: back cover -- -- `Chartar Gyugh' 1: 5 -- -- `Fontenay Quince' 1: 5 -- -- `Harron' 1: 6 -- -- `Orange', 1922 illustration 1: 4 -- -- `Pineapple' 1: 2, 2, 4 -- -- `Smyrna' 1: 4 -- -- `Van Deman' 1: 2 -- sinensis 1: 8, 8 \"Cydonia oblonga: The Unappreciated Quince,\" Joseph Postman 1: 29, 28, back cover Cytokinin pathways 4: 1519 D Da Pao Shan mountain 3: 12, 13 -- -- -- climate 3: 8, 9, 13 Darwin, Charles 4: 16, 26 -- -- The Power of Movement in Plants 4: 16 Darwin, Francis 4: 16 Dating of trees 2: 3031 David, Father Armand, discoveries revisited 2: 1, 2228 -- biography 2: 24 Index 37 -- portrait 2: 24 Davidia 2: 22 -- involucrata, in China 2: 24, 25, 25 Dawn redwood 4: 23 Da Xue Shan Mts. 3: 4, 5, 89, 1213 Daylilies, shade-grown 3: 33 Deforestation 1: 21 Del Tredici, Peter, \"The Sand Pear-- Pyrus pyrifolia\" 4: 28 -- -- -- photographs by 1: inside back cover; 4: inside covers Den Boer, Arie F. 2: 2 Dengchigow mountain 2: 28 Deutzia glomeruliflora, in China 2: 26 Dipelta yunnanensis, in China 2: 25 Dirr, Michael 2: 32; 3: 21 Diseases of pome fruits 1: 4, 6, 7, 8 Disjunct flora 1: 44 Disporum bodinieri, in China 2: 25 Diversification, in black oaks 4: 413 Diversity in alpine habitat 3: 6, 1113 Dogwood, in China 2: 25, 26 -- red-twig, as woody floral 3: 33, 34 Dormancy, and tree hormones 4: 17 Dosmann, Michael S., \"Autumn's Harbinger: Acer rubrum `Schlesingeri'\" 2: 32, inside back cover -- -- -- \"Malus at the Arnold Arboretum: An Ongoing Legacy\" 2: 1421 Douglas firs, as beetle host 1: 33, 35 Drepanostachyum, in China 2: 26 Dulce de membrillo 1: 45 Erwinia amylovora 1: 6; 2: 10; 4: 22 \"Essay on Naming Nature: The Clash Between Instinct and Science,\" P.F. Stevens [review] 4: 2527 Ethanol, in beetle trap 1: 32 Ethylene 4: 15 Euonymus alata, restrictions on 3: 23 -- in China 2: 28 Eurasia, quinces in 1: 47 Eurasian beetles, damaging 1: 3335 Europe, quinces in 1: 45, 7 European spruce bark beetle 1: 35 Evolutionary relationships, and oaks 4: 13 -- -- and classification 4: 2527 \"Excerpt From Wilson's China: A Century On,\" Mark Flanagan and Tony Kirkham 3: inside front cover, 213, 36, 812 Exotic beetles 1: 3135 Experiential learning 3: 32 \"Extraordinary Discoverer of Life\" 2: 24 -- health checklist 3: 27 -- non-timber crops from 3: 2635 -- private ownership of 3: 2728 \"Forest Farming,\" Ken Mudge 3: 2635, 2634 Forest Farming Practicum [Cornell] 3: 32 Forest Hills Station 1: 12 Forests, North American oak 4: 213 Franchet (the botanist) 2: 27 Fraxinus, and exotic beetles 1: 34 Fruit, nutraceutical 3: 1425 -- spot 1: 6, 7 -- trees 1: 29; 4: 28 Fuji cherry 4: 24 Fuller, Dave 1: 19 Fullerview Photography 1: 19 Fumariaceae 2: 26 Fungus 1: 33, 35; 2: 10, 20; 4: 20 G \"Golden Apple\" of antiquity 1: 23 Garden and Forest 1: 44 Genealogical relationships 4: 2527 Genebank, national system 1: 6, 7, 9 Gene flow, between oak spp. 4: 711 -- -- interspecific 4: 9 Genera Plantarum [Bentham and Hooker] 4: 26 Genetics, in Aronia 3: 19, 21, 22, 2425 -- in Malus 2: 20 -- nut-tree grafts and 3: 32 -- of black oak group 4: 213 Geographic distance, and oak speciation 4: 213 -- information systems (GIS) 1: 10, 19 Germplasm Resources Information Network (GRIN) 3: 21 Gibberellins 4: 15 Ginseng, American, 3: 2830, 29, 30 -- -- growing 3: 2830, 32 -- -- market for 3: 2930 -- -- medicinal uses 3: 2930 -- -- pricing 3: 30 -- -- root 3: 30 -- -- soil calcium and 3: 29 Ginsenosides 3: 30 Glaciation, and oaks 4: 13 Gleason, Herbert Wendell 1: 13 Globalization 2: 29 F Fabraea maculata, and quince 1: 6, 7 Fagus and exotic beetles 1: 35 Fairchild Aerial Surveys, Inc. 1: 10, 11, 18 Fairchild, Sherman Mills 1: 10 Famiglietti, Bob, \"2009 Weather at the Arboretum\" 4: 2024 \"Farming or Wildcrafting?\" 3: 27 Fengtong nature reserve 2: 22, 22, 2428, 25, 26 Ferns, shade-grown 3: 33 Fertilizing, and tree hormones 4: 18 Fiala, Father John 2: 6, 14 Field-forest ecotone 3: 28 Fir, Nikko 4: 22 -- as beetle host 1: 33, 35 Fire blight 1: 6, 7, 8; 2: 10 -- and oaks 4: 2 Flanagan, Mark 3: 2, 3 -- -- \"An Excerpt From Wilson's China: A Century On,\" with Tony Kirkham 3: 213 Forest ecology 2: 2931; 3: 2628 -- farming, introduction to 3: 2635 -- -- crop selection for 3: 2830, 33, 35 -- -- income potential of 3: 26, 28, 29, 30, 33, 35 E \"Early Bloomer: Hydrangea paniculata `Praecox',\" Sue A. Pfeiffer 1: inside front\/back covers, 44 Eastern Aerial Surveys, Inc. 1: 15 East Malling quince rootstocks 1: 5 Ecology, and oaks 4: 13 -- of forests 2: 2931; 3: 2628 Edgar, Mr. [circa 1910] 3: 4, 6, 7 Edible crops, new 3: 1425, 3033 Education, experiential 3: 32 Egolf, Don 2: 7 Elms, as beetle host 1: 34 Emerald ash borer 1: 34, 34 Enkianthus deflexus, in China 2: 25, 25 Epimedium davidii, in China 2: 26, 27 Ericaceae 2: 24, 25 38 Arnoldia 68\/1 Goldenseal 3: 29, 29 Gongga Shan Mt. (Minya Konka) 3: 4, 5, 810 Google Earth 1: 10 GPI Models 1: 18 GPS database, in curriculum 3: 32 Gravitropism 4: 16 Gray, Asa, and disjunct flora 1: 44 Great Lakes region, oaks in 4: 213, map 6 Griffola frondosa 3: 30 Growth processes, and plant hormones 4: 1519 -- rate, measuring 2: 3031 Grunsfield, John 1: 16 H Hardiness, of camellias 1: 2030 -- of crabapples 2: 3 -- of hydrangea 1: 44 -- of pines 3: 36 Hardwoods, for mushroom-growing 3: 31 -- secondary 3: 32 Harvard Institute for Geographical Exploration 1: 12, 16 Harvard Real Estate 1: 17 Hayrapetyan, Vagharshak, Dr. 1: 4 Hellebore, Tibetan 2: 2728, 28 Hellebores, shade-grown 3: 33 Helleborus chinensis 2: 2728 -- thibetanus, in China 2: 2728, 28 Hemlocks, as beetle host 1: 35 Hengduan Shan mountains 3: 413 Hen-of-the-woods mushroom 3: 30 Hericium spp. 3: 30, 31 Heucheras, shade-grown 3: 33 Hickory nuts 3: 30, 32 -- red pignut 3: 32 -- shagbark 3: 32 -- varieties and grafts 3: 32 Hill, Rev. Ernest J. Hill 2: 4 \"Hill's Oak: The Taxonomy and Dynamics of a Western Great Lakes Endemic,\" Andrew L. Hipp 4: 213, 26, map 6, genetic chart 7, 812 Himachal Pradesh, pine from 3: 36 Himalayan National Park 3: 36 Himalayan pine 3: 36, inside back cover Himalayas, Eastern 3: 3, 413, 5, 8, 9, 12 -- flora of 3: 213, 36 Hines Nursery 1: 28 Hipp, Andrew L. \"Hill's Oak: The Taxonomy and Dynamics of a Western Great Lakes Endemic,\" 4: 213 -- -- -- photo by 4: back cover Hippeastrum 4: 16 Hokkaido 1: 44 Holboellia, in China 2: 26, 27 Holden Arboretum 1: 21 Honeysuckle family 2: 25 Honeysuckles, in China 2: 28 Honshu 1: 44 Hood Blimp 1: 19 Hooker, Joseph Dalton 4: 26 Hormones, and plant care 4: 1519 \"Hormones and Tropisms\" 4: 16 Hornbeam 2: 26; 3: 31 Horsechestnut, as beetle host 1: 34 Hosie, Sir Alexander 3: 11 Hostas, forest-grown 3: 33, 33 Howard, Heman 1: 18 -- -- 1959 photo by 2: 17 Hubble telescope camera 1: 16 Hubei Province 2: 19; 4: 28 Humanity and trees 2: 2931 Hun, Chang Yong 1: 21 Hybrid, intergeneric 1: 34; 3: 25 -- interspecific 3: 21 Hybrization, in oaks 4: 213 Hydrangea, panicle 1: 44 Hydrangea paniculata `Praecox' 1: inside front\/back covers, 44 Hydrastis canadensis 3: 29, 29 Hylurgus ligniperda 1: 33, 33 Iowa State University 2: 13 Ips sexdentatus 1: 33, 33 -- typographus 1: 35 -- -- spruces killed by 1: 35 Iran, quince germplasm in 1: 7 Ironwood 3: 31 Irrigation, and hormones 4: 1819, 18 Japan, plants from 1: 30, 44; 2: 16 Japanese spicebush 1: 24 Johnson, Ethan W. 1: 15 Joyce Kilmer Park, in aerial photos 1: 1936, 14; 1955, 18 Juglans spp. 3: 30 Juice crops, sustainable 3: 1425 June, Chang Yong 1: 21 Junipers, alpine 3: 8 Kabul, pine from 3: 36 Kangding expedition 3: 213, 10 Kazakhstan wild apple 2: 20 Kelley, Susan 1: 17 Kenya, agroforestry center 3: 27 Kew's Arboretum 3: 2 -- expeditions to Himalayas 3: 213, 36 Kilimanjaro, agroforestry on 3: 27 Kirkham, Tony 3: 2, 3 -- -- \"An Excerpt From Wilson's China: A Century On,\" with Mark Flanagan 3: 213 -- -- photo by 3: inside front cover Knox, Charles 1: 2 Koller, Gary 2: 32 Korean camellias 1: 2030, 23, 2530 -- climate 1: 2025, 30 -- mountain ash 1: 24 Kuan Hsien, travelling in 3: 1112 IJ K Iles, Jeff, \"Crabapples...With No Apologies\" 2: 213 Illinois, oaks in 4: 213 Incarvillea delavayi 3: 13 -- in a Tibetan scene 3: 11 \"Index to Arnoldia, Volume 66\" 1: 3643 India, pine from 3: 36 Indiana, spruce beetle in 1: 35 International Council for Research in Agroforestry [Kenya] 3: 27 \"In the Footsteps of Father David,\" Cedric Basset 2: 2228, 2228 Iowa, Aronia in 3: 32 L Ladyslipper, Tibetan 3: 12 Lampshade poppy, quest for 3: 2, 47 Landscape ornamentals 1: 29, 2030, 44; 2: 221, 32; 3: 1425, 36; 4: 28 Land-use 3: 2635 Larches, as beetle host 1: 33, 35 Lardizabalaceae 2: 26 Larix spp., and exotic beetles 1: 33, 35 Latin America, quinces in 1: 45 Leaf-spot 1: 6, 7 -- frogeye 2: 10 Leaves, soil nutrients in 3: 29 Lentinula edodes 3: 28, 30 Index 39 Leptographium spp. 1: 33 Lian Lua Shan mountains [Tibet] 3: 5 Lightning damage 4: 22 Lilium lophophorum 3: 13 Lindera obtusiloba, autumn foliage 1: 24 Lindgren funnel trap 1: 32, 33, 35 Linnaean system and new trends 4: 2627 Linnaeus, Carolus, statue of 4: 27 Lion's mane mushroom 3: 30, 31 Liriodendron tulipifera 3: 31 Lhasa, travelling to 3: 25 Longwood Gardens, camellias at 1: 28 Lonicera, in China 2: 28 Lyon Botanical Garden 2: 28 M MacDaniels, Lawrence 3: 32 \"MacDaniels Nut Grove: A Unique Educational Site\" 3: 32, 32 Magnolia, in China 2: 28 -- liliiflora, in China 2: 23 Maine, chokeberry in 3: 19 -- exotic beetle in 1: 35 Maitake mushroom 3: 30 Malus 1: 3; 2: front\/back covers, 221 -- `Adirondack' 2: 7, 7, 9 -- `Aldenhamensis' 2: 2 -- `Almey' 2: 2 -- `Amberina' 2: 5 -- `Antonovka' rootstock 2: 18 -- apple-scab and 2: 3, 10, 10 -- Arboretum legacy of 2: 1421, back cover -- x atrosanguinea 2: 17 -- baccata 2: 5, 1920 -- -- `Columnaris' 2: 21 -- -- `Jackii' 2: 21 -- `Barbara Ann' 2: 21 -- bark interest 2: 1920 -- `Blanche Ames' 2: 17, 21 -- `Bob White' 2: 10, 10, 21 -- `Camzam' (Camelot) 2: 7, 8, 9 -- `Cardinal' 2: 7, 9 -- centenarian 2: 16 -- `Cinzam' (Cinderella) 2: 7, 9 -- `Coralcole' (Coralburst) 2: 7, 7 -- `David' 2: 7, 8, 9 -- diseases and resistance to 2: 3, 6, 910, 20 -- documentation 2: 16 -- `Dolgo' 2: 10, 10 -- `Donald Wyman' 2: 8, 9, 10, 11, 19, 19, 21 -- `Dorothea' 2: 2, 21, 21 -- `Doubloons' 2: 13 -- dwarf 2: 7, 18 -- floribunda 2: 2, 4, 6, 9, 10, 16 -- -- `Exzellenz Thiel' 2: 6 -- -- Selection #821 2: 20 -- flowering display 2: 2, 24, 4, 68, 9, 1219, 13, 17, 19, 21 -- foliage 2: 3, 5, 910 -- fruiting 2: front\/back covers, 3, 4, 5, 56, 8, 910, 10, 11, 13, 19, 19, 20 -- -- preserves from 2: 10 -- fruitless cultivars 2: 10 -- genetics and selection 2: 1417, 20 -- halliana 2: 20 -- hardiness 2: 3 -- `Henrietta Crosby' 2: 17, 21 -- `Henry F. Dupont' 2: 17, 21 -- `Hopa' 2: 3 -- `Huber' (Royal Fountain) 2: 13 -- hupehensis 2: 16, 17, 20 -- `Indian Magic' 2: front cover, 10 -- ioensis `Palmeri' 2: 21 -- `Jarmin' (Marilee) 2: 10 -- `Jewelcole' (Red JewelTM) 2: 8, 9, 10 -- `JFS-KW5' (Royal Raindrops) 2: 9, 13 -- `Katherine' 2: 21 -- lancifolia hybrids 2: 18 -- `Lanzam' (Lancelot) 2: 7, 9 -- `Liset' 2: 2, 2, 6 -- `Lollizam' (Lollipop) 2: 7 -- longevity 2: 16 -- `Louisa' 2: 9, 13, 13 -- `Mary Potter' 2: 17, 17, 21 -- `Orange Crush' 2: 6, 6, 9 -- pedicel variation 2: 19 -- `Pink Pearl' 2: 21 -- `Prairie Maid' 2: 5, 9 -- `Prairifire' 2: 9, 10, 12, 13 -- `Prince Georges' 2: 21 -- `Profusion' 2: 2 -- prunifolia 2: 56 -- -- `Pendula' 2: 6 -- `Purple Prince' 2: 7, 9 -- `Radiant' 2: 3 -- `Red Jade' 2: 2, 5, 56, 12, 13 -- `Red Swan' 2: 5, 6 -- x robusta `Erecta' 2: 21 -- rootstock effects 2: 20 -- sargentii 2: 6, 16 -- -- `Rosea' 2: 7, 17, 21 -- -- `Select A' (Firebird) 2: 7 -- -- `Tina' 2: 7 -- `Satin Cloud' 2: 5 -- Sax experimental hybrids 2: 18, 18 -- `Schmidtcutleaf' (Golden Raindrops) 2: 7, 8, 9 -- sieversii 2: 20 -- `Snowdrift' 2: 10, 11 -- `Spring Snow' 2: 10 -- sylvestris hybrids 2: 18 -- transitoria, in China 2: 15 -- tschonoskii 2: 5, 16 -- unnamed #691-52-A 2: 18 -- variation of 2: 3 -- wasps and 2: 3 -- weeping forms 2: 56, 1213, 13 -- winter interest 2: 11, 12, 13, 20 -- x zumi `Calocarpa' 2: 21 \"Malus at the Arnold Arboretum: An Ongoing Legacy,\" Michael S. Dosmann 2: 1421, 1421 \"Malus Mystery\" 2: 18 Maple 3: 32, 36 -- David's 2: inside front cover, 22 -- products 3: 28, 35 -- red, cultivar `Schlesingeri'2: 32, inside back cover -- red 3: 31 -- sugar 3: 31 Maples, as beetle host 1: 35 Maps from photos 1: 10, 1215, 1719 March, Sylvester 1: 21 Marino, Sergio 1: 18 Maryland, spruce beetle in 1: 35 Maslow, Abraham, human needs concept 2: 29 Massachusetts Department of Conservation and Recreation 1: 34 Massachusetts forests 3: 26 Massachusetts Public Works 1: 18 Maximowicz (the botanist) 2: 28 Mayr, Ernst 4: 25, 26 Meconopsis x beamishii 3: 7 -- x finlayorum 3: 7 -- horridula 3: 13 40 Arnoldia 68\/1 -- integrifolia ssp. integrifolia, in China 3: inside front cover, 2, 47, 4 -- -- x grandis 3: 7 -- -- x quintuplinervia 3: 7 -- pseudointegrifolia 3: 7 Medicinal crops 3: 2930 Mediterranean beetles, damaging 1: 33 Mesopotamia, quince in 1: 3 Mespilus 1: 4; 3: 21 Metasequoia glyptostroboides, fall color 4: 23 Mexico, oaks native to 4: 3 Meyer, Paul 1: 21, 22, 23, 24 Michigan, EAB in 1: 34 Microbiota decussata cutting 4: 18 Midwest, Aronia in 3: 23 Miller, George 4: 26 Min Shan mountains 2: 1 Missouri Botanical Garden 4: 27 Morchella spp. 3: 30 Morels 3: 27, 30 Morphology, oak speciation and 4: 213 -- scientific classification and 4: 2527 Morris Arboretum, camellia selection at 1: 2030, 2530 -- -- -- Korean expeditions of [19791991] 1: 2024, 2023 Morton Arboretum, oak research at 4: 414 -- -- herbarium specimens 4: 35, 9 Mountain ash, Korean 1: 24 -- habitats 2: 2228 -- -- Afganistan 3: 36 -- -- Burma 3: 36 -- -- Maine 3: 19 -- -- Tennessee 3: 20 -- -- Sichuan\/Tibet 3: 213, 3, 5, 8, 9, 12 Mt. Emei 2: 26 Mt. Wachusett 3: 26 Mudge, Ken, \"Forest Farming\" 3: 2635 Murray, Joseph, \"Tree Hormones and Why They Matter\" 4: 1519 Museum of Science 1: 16 Mushroom cultivation 3: 28, 3033, 31, 33 -- income from 3: 30 -- nutrient process 3: 31 -- spawn 3: 31, 33 -- wild-collected 3: 27 Mutagen breeding 3: 24 N Nadkarni, Nalini M., Dr. 2: 29 -- -- -- \"Between Earth and Sky: Our Intimate Connections to Trees\" [excerpt] 2: 2931 Nakai (Japanese botanist) 4: 28 Naming Nature: The Clash Between Instinct and Science [Yoon, reviewed] 4: 2527 National Arboretum 2: 7 National Plant Germplasm System (NPGS) 1: 6, 7, 9 Native Americans and plants 3: 27, 29 Nebraska, Aronia in 3: 23 Nematode, pinewood 2: 29 New England, hydrangeas in 1: 44 Newton, Amanda A., 1909 illustration by 1: back cover New York forests 3: 2728 Nightshade family, blights 4: 20 Nikko fir 4: 22 Nineteenth-century aerial photography 1: 10, 10 -- -- garden trends 1: 44 Nomenclature, binomial 4: 26 -- of Aronia 3: 21 Non-native pests 1: 3135 North America, flora of 1: 44; 3: 1425 -- -- forest-farming in 3: 27 -- -- oaks of 4: 213, map 6 -- -- pests from 2: 29 North Carolina, chokeberry in 3: 18 Northeast Aerial Photos 1: 15 Northeast, versatile shrub for 3: 1425 -- -- reforestation trends in 3: 26, 28 Nuclear data, and black oaks 4: 3 Nursery trade 2: 14, 32 Nut groves 3: 30, 32 Nutraceutical fruit crop 3: 1425 -- -- petioles 4: 11 -- genome 4: 711 -- Hill's, disturbance and 4: 13 -- -- interbreeding and taxonomy 4: 113, 25, map 6, genetic chart 7, 910 -- northern pin 4: 213 -- pin 4: 5 -- red 4: front cover -- scarlet 4: 310, 4, 6, map 6, genetic chart 7, 89, 12 -- white 4: 12 Oaks, as beetle host 1: 35 -- black group 4: 213, map 6, genetic chart 7 -- white group 4: 23 Ohio Valley, camellias in 1: 27 Olmsted style 1: 6, 12 Ophiostoma spp. 1: 33 Orchids, wild 2: 25 Oregon, Aronia in 3: 23 -- exotic beetle in 1: 35 -- USDA genebank in 1: 7, 9 Ornamentals, forest-grown 3: 3335, 33, 34 -- woody 1: 29, 2030, 44; 2: 221, 32; 3: 1425, 36; 4: 28 Osmanthus forest, in China 2: 24 Oyster mushroom 3: 30, 31 P Panax quinquefolius, as crop 3: 2830, 29, 30 Panda, giant 2: 22, 23, 24, 28 Pao-chung, Kao, Prof. 2: 28 Paris fargesii, in China 2: 25 Parks, Clifford, Dr. 1: 24, 28 Pawpaw fruit 3: 28, 28, 30 Pear 1: 3, 4, 5 -- `Beurre Hardy' 1: 5 -- `Bradford' Callery 4: 28 -- `Comice' 1: 5 -- Japanese 1: 4 -- `Old Home' 1: 5 -- quince hybrid 1: 4, 4 -- quinces and 1: 37 -- sand, at Arboretum 4: inside covers, 28 -- -- fruit of 4: 28 -- -- vigor of 4: 28 -- tribe 1: 3 O Oak 3: 31, 32, 36 -- black 4: 56, 1013, 1011, back cover -- -- hybrids 4: 7 Index 41 Pectin 1: 2; 3: 23 Perennials, pot-in-pot 3: 33, 33 -- shade-grown 3: 33 Permaculture 3: 32 Pests 1: 3135 Pfeiffer, Sue A., \"Early Bloomer: Hydrangea paniculata `Praecox'\" 1: inside front\/back covers, 44 Pheneticists, and trends 4: 26 Philadelphia, camellias for 1: 20, 2430 Photinia 3: 21 -- floribunda 3: 21 -- melanocarpa 3: 21 -- pyrifolia 3: 21 Photography, aerial 1: 1019 Photosynthesis and plant hormones 4: 1819 Phototropism 4: 16, 16 Phytophthora infestans 4: 20 Picea spp., and beetle damage 1: 33, 35 -- forest in Slovakia 1: 35 Pine bark beetle, red-haired 1: 33, 33 -- cones for crafts 3: 27, 33 Pine, Eastern white 3: 36 -- Himalayan 3: 36, inside back cover -- lanceolate 2: 22, 23 -- Monterey, with bark damage 1: 33 Pines, as beetle host 1: 33, 35 -- cold hardiness of 3: 36 Pinewood nematode, in Europe 2: 29 Pinus spp., and exotic beetles 1: 33, 35 -- excelsea 3: 36 -- griffithii 3: 36 -- nepalensis 3: 36 -- radiata, with beetle damage 1: 33 -- strobus 3: 36 -- thunbergii 1: 24 -- wallichiana 3: 36, inside back cover -- -- synonyms for 3: 36 Plant exploring 1: 2030; 2: 2228; 3: 213 -- hormones, and arboriculture 4: 1519 Plant Protection and Quarantine (PPQ) program 1: 3135 Pleione limprichtii, in China 2: 25 Pleurotis spp. 3: 30, 31 Ploidy, in Aronia 3: 1922 Plums 2: 28 Polly Hill Arboretum 1: 27 Pome fruits 1: 27 Poppies, lampshade 3: inside front cover, 2, 47, 4 Postman, Joseph, \"Cydonia oblonga: The Unappreciated Quince\" 1: 29 Powdery mildew 1: 6 Powerline undergrowth 3: 18 Practicum in Forest Farming (Cornell course) 3: 32 Primula secundiflora 3: 6 Pruning, and hormones 4: 1519, 17, 19 Prunus spp., in China 2: 28 -- incisa f. serrata 4: 24 Pseudocydonia sinensis, in North America 1: 8, 8 -- -- flowers and bark 1: 8 Pseudotsuga spp., and exotic beetles 1: 33, 35 Psychology, and trees 2: 29, 30 Pujigou 2: 22, 2425 Purdue-Rutgers-Illinois Apple Breeding program 2: 20 Pyreae 1: 3 Pyronia 1: 4, 4 -- veitchii 1: 4 Pyrus calleryana `Bradford' 4: 28 -- pyrifolia 1: 4; 3: 21; 4: inside covers, 28 -- serotina 4: 28 -- sinensis 4: 28 Q Quarryhill Botanic Garden 3: 36 Quercus spp., 3: 31 -- -- exotic beetles and 1: 35 -- -- genetic studies of 4: 513 -- coccinea 4: 1, 310, 4, map 6, genetic chart 7, 89, 12 -- -- acorns 4: 4 -- ellipsoidalis, related spp. and 4: 113, 25, map 6, genetic chart 7, 910 -- -- acorns 4: 3, 5, 10 -- imbricaria 4: 3 -- Lobatae Section 4: 3 -- x palaeolithicola 4: 7 -- palustris 4: 3 -- phellos 4: 3 -- pumila 4: 3 -- rubra 4: front cover, 3 -- shumardii 4: 3 -- velutina 4: 313, 56, genetic chart 7, 1011, back cover -- -- acorns 4: 5, 1011 Quince 1: 29, 28, back cover -- \"A\" rootstock clone 1: 5 -- `Angers' 1: 5 -- \"C\" rootstock clone 1: 5 -- `Champion', 1909 illustration 1: back cover -- `Chartar Gyugh' 1: 5 -- Chinese 1: 8, 8 -- cold-hardiness 1: 6, 7, 8 -- culinary uses 1: 2, 45 -- cultivars 1: 27 -- diseases or problems 1: 4, 67, 8 -- flowers and foliage 1: 3 `Fontenay' 1: 5 -- fruit 1: 2, 2, 47, 47. back cover -- genetics 1: 4, 5, 67, 9 -- `Harron' 1: 6 -- history of 1: 26 -- hybrids 1: 4 -- in Kakheti [Eurasia] 1: 7 -- landscape qualities 1: 2, 56, 8 -- `Orange', 1922 illustration 1: 4 -- pear hybrid 1: 4 -- pears and 1: 37 -- pectin content 1: 2 -- `Pineapple' 1: 2, 2, 4 -- propagation and grafting 1: 27 -- Provence 1: 5 -- pruning 1: 6 -- rootstocks 1: 3, 4, 5, 6, 7 -- Shilda variety 1: 7 -- `Smyrna' 1: 4 -- `Van Deman' 1: 2 R Radiation, and plant breeding 3: 24 Radioactive \"bookmark\" of 1954, and tree-dating 2: 31 Rainforest, dating trees in 2: 3031 -- experience 2: 30 Ramps 3: 30 Ranunculaceae 2: 27 Raspberry, black 3: 28 42 Arnoldia 68\/1 Red osier dogwood 3: 34 -- haired pine bark beetle 1: 33, 33 Reed, Dr. George M. 2: 6 Reeve, Bob, archival photograph by 1: 16 Reforestation trends 3: 28 Rehder, Alfred, 1: 44; 4: 28 -- Malus and 2: 1617 Resistance breakdown, in crabapples 2: 10 Rheum alexandre 3: 6 Rhododendron, in Sichuan 2: 22, 22 -- capitatum 3: 13 -- prezwalskii, in Tibet 3: 10 Rhododendrons, alpine 3: 8, 10, 13 Rock, Joseph F., and Gongga Shan 3: 4 -- -- 1926 photo by 2: 15 Root development, and hormones 4: 1519, 18 -- disease fungi 1: 33 Rosaceae 1: 34, 6; 2: 16; 3: 14, 19 Rose, Nancy, \"A Soft Touch: Pinus wallichiana\" 3: 36, inside back cover -- -- photos by: 2: front\/inside back covers; 3: front\/inside back covers; 4: front cover Royal Botanic Gardens, Kew 3: 2, 36 Rubus spp. 3: 28 Russia, Aronia breeding in 3: 22, 23, 24, 25 Rust diseases 1: 6 Rutgers, apple breeding at 2: 20 S Sakhalin Island 1: 44 \"Sand Pear--Pyrus pyrifolia,\" Peter Del Tredici 4: inside covers, 28 Sapindaceae 2: 26 Sargent, Charles S. 2: 14, 16, 17, 19, 32 -- -- -- Japanese plants and 1: 44 -- -- -- photography and 3: 1011 -- -- -- Wilson, E.H. and 3: 2, 1011 Sasaki Associates 1: 17 Sax, Karl 3: 36 -- -- Malus research and 2: 1719 Schlesinger, Barthold 2: 32 Science in the Pleasure Ground exhibit 1: 1819 Scientific Center for Viticulture [Armenia] 1: 5 Scientific classification 4: 2527 -- limits 4: 27 Scott Arboretum 1: 27 \"Searching for Exotic Beetles,\" Nichole K. Campbell 1: 3135, 3135 Seasonal changes, and trees 2: 30; 4: 1517, 19 Seed collection 1: 20, 2324 \"Seeking Cold-Hardy Camellias,\" Anthony S. Aiello 1: 2030, 2030 Shaw, Peter Ashton 1: 15 Shawnee National Forest 4: 8, 12 Shiitake cultivation 3: 28, 3033, 31, 33 Shipping industry 1: 3132; 2: 29 Shrubs, centenarian or notable specimens 1: 44, inside back cover -- native 3: 1425 Sichuan\/Tibet 2: 2228; 3: 213, 3, 5, 8, 9, 12 Sino-American Botanical Expedition [1980] 2: 19 Sinofranchetia chinensis 2: 26 Six-toothed bark beetle 1: 33, 33 Slovakia, bark beetle in 1: 35 Sochong Island 1: 22, 23 \"Soft Touch: Pinus wallichiana,\" Nancy Rose 3: 36, inside back cover Soil 3: 29 -- and oaks 4: 13 Solanaceae, blights of 4: 20 Sorbus 1: 4; 2: 28; 3: 21 -- alnifolia 1: 24 -- aucuparia and Aronia cross 3: 25 South Korea, deforestation in 1: 21 -- -- expeditions to 1: 2024, [1984] map 21, 2023, 30 -- -- hardy plants from 1: 2030 -- -- landscape 1: 2123 Spath Nursery [Berlin] 2: 6, 32 Species, biological concept 4: 2, 25 -- differentiation in black oaks 4: 213 Spicebush, Japanese 1: 24 Spiraeoideae 1: 3 Spruce bark beetle 1: 35 Spruces, as beetle host 1: 33, 35 Stevens, P.F., \"An Essay on Naming Nature: The Clash Between Instinct and Science\" [review] 4: 2527 Stewartia pseudocamellia, branching 4: 15 Storm damage 4: 22 -- -- plant hormones and 4: 19 Street trees, beetle damage to 1: 34 Strontium, radioactive 2: 31 Styrax japonica 1: 24 Sunlight, and plant hormones 4: 16 Sustainable crops 3: 23, 2635 Swissair Photos + Surveys 1: 15 Sympatry, among oaks 4: 3 Systematist debates 4: 2527 T Tachien-lu 3: 2, 8, 11 Taean Peninsula 1: 21 Taechong Island 1: 2023 Tagong temple 3: 12, 13 Talltree Arboretum 4: 9 Tang li tzu 4: 28 \"Taxonomic Teasers in Aronia\" 3: 21 Taxonomy, of Aronia 3: 21 -- -- black oaks 4: 213 -- -- Malus 2: 14, 1617, 20 -- trends in 4: 2527 Three-ips lure, in beetle trap 1: 32 Thuja spp., and exotic beetles 1: 35 Tibetan Empire 3: 23 Tibetan Frontier, circa 1910 3: 213, 8 Tibetan region, exploration in 2: 2228; 3: 213 -- peoples 3: 23, 4, 11 Time concepts, and trees 2: 29, 3031 Tinley Creek Forest Preserve 4: 10 Topping damage 4: 19, 19 Toucans 4: 25 Tournachon, Gaspard-Felix (Nadar) 1: 10 Trabut, Louis 1: 4 Trapping beetles, chemistry of 1: 32 Tree care, and hormones 4: 1519 -- consumption 2: 29 -- dating 2: 3031 -- longevity 4: 15 Index 43 \"Tree Hormones and Why They Matter,\" Joseph Murray 4: 1519, 1519 Trees, benefits of 2: 2930; 4: 15 -- centenarian or notable specimens 2: 16, 19, 32, inside back cover; 3: 36, inside back cover; 4: 22, 22, 23, 28, inside back cover -- damage to 4: 18, 18, 19, 22, 24 -- human needs met by 2: 2931 -- radioisotopes and dating of 2: 31 -- seasons and 2: 30; 4: 1517, 19 -- time and 2: 20, 29, 3031 -- urban 4: 1519 Trilliums, shade-grown 3: 33 Tropical forests 2: 3031; 3: 27 Truffles, difficulty of 3: 30 Tsuga spp., and exotic beetles 1: 35 Tuber spp. 3: 30 Tulip poplar 3: 31 Turkey, quinces in: 1: 4, 6, 7 Turkmenistan, fruit trees in 1: 4, 7 Tyler Arboretum 1: 27 UV W \"Umwelt\" concept 4: 2627 Urban mapping 1: 17 -- tree care 4: 1519 United States Department of Agriculture (USDA) 1: 2 -- -- -- -- APHIS and beetles 1: 3135 -- -- -- -- artwork, circa 1900 1: back cover -- -- -- -- GRIN 3: 21 -- -- -- -- hardiness zone five 3: 36 -- -- -- -- links 1: 34; 3: 21 -- -- -- -- NPGS 1: 6, 7, 9 -- -- -- -- PPQ program 1: 3135 -- -- -- -- Plants Database 3: 21 -- -- -- -- Trans-Caucasus expeditions, recent 1: 4, 7 US economy and tree consumption 2: 29 US Geological Surveys (USGS) 1: 17 US National Agricultural Library, 1909 illustration from 1: back cover US National Arboretum 1: 8, 21 -- -- -- -- winters at 1: 24 University of Bristol, Long Ashton [Eng.] 3: 21 University of Connecticut, Aronia research at 3: 1425 University of Illinois, apple breeding at 2: 20 University of North Carolina, Chapel Hill 1: 24 University of Pennsylvania arboretum 1: 30 University of Wisconsin, juice crop research 3: 23 Vaccinium corymbosum 3: 23 Vavilov Institutes [Soviet] 1: 7 Veitch Nurseries, E. H. Wilson and 3: 2, 10 Venturia inaequalis 2: 10 Vf gene, and apple scab 2: 20 Viburnum bitchuense 1: 24 -- brevitubum, in China 2: 26, 27 -- chingii 2: 26 Viburnums 2: 26 Wachusett, view from 3: 26 Walnut 3: 30, 32 -- varieties and grafts 3: 32 War, deforestation from 1: 21 Warming trends 4: 24 Washburn, Bradford 1: 16 -- -- aerial photography of 1: 1217, 13, 14 Wasp, yellow-jacket 2: 3 Watersprouts 4: 19 Watson, James 4: 26 \"Weather at the Arboretum--2009,\" Bob Famiglietti 4: 2024, chart 21, 2224 Weather damage at Arboretum 4: 22 Weather Station Data, 2009 4: 21 Wildcrafting 3: 27, 33 Willow, corkscrew 3: 35 Willows, as beetle host 1: 34 Willowwood Arboretum 1: 27 Wilson, Ernest Henry, hydrangea and 1: 44 -- -- -- lampshade poppy and 3: 2, 47 -- -- -- Malus finds 2: 16 -- -- -- sand pear and 4: 28 -- -- -- Tibetan discoveries revisited 3: 213 Wilson's China: A Century On [Flanagan and Kirkham, excerpt] 3: 213 Windsor Great Park 3: 2 Wisconsin, ginseng cultivation in 3: 30 Wood beetles 1: 3135 -- digestion by mushrooms 3: 31 -- industry 2: 29 -- -- pests and 1: 3135 -- packing as pest vector 1: 3132, 34; 2; 29 Woody florals 3: 3335 -- ornamentals 1: 29, 2030, 44; 2: 221, 32; 3: 1425, 33, 35, 36; 4: 28 Worcester, MA, ALB in 1: 34 World Agroforestry Center [Kenya] 3: 27 World War II, deforestation from 1: 21 -- -- -- nursery trade and 2: 14, 32; 3: 3335 Wright, Wilbur 1: 10 Wyman, Donald, aerial photography of 1: 18 -- -- crabapple namesake 2: 8, 9, 11, 19, 19, 21 -- -- -- legacy 2: 19 -- -- quince opinion 1: 56 -- -- `Schlesingeri' red maple and 2: 32 XYZ Xizang Autonomous Region 3: 3 Xyleborus spp. 1: 35 -- seriatus 1: 35 Xylotrechus spp. 1: 35 -- hircus 1: 35 Ya-jia Pass region 3: 310, 3, 6, 8, 9 Yangtze River, Wilson in 3: 11 Yellow Sea islands 1: 2021, 2123 Yichang, Wilson in 3: 11 Yinger, Barry 1: 20, 21, 22, 28 Yoon, Carol Kaesuk, at Cornell 4: 25 -- -- -- Naming Nature: The Clash Between Instinct and Science [reviewed] 4: 2527 Yushania, in China 2: 26 Zacharias, Elizabeth H., Ph.D. 2: 28 Zeiss camera on Hubble 1: 16 Zhong, Xiao 3: 13 Zhedou Pass 3: 45, 12 Compiled by Rosalie Davis "},{"has_event_date":0,"type":"arnoldia","title":"A New Plant Introduction from the Arnold Arboretum: Ilex glabra 'Peggy's Cove'","article_sequence":5,"start_page":44,"end_page":45,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25480","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060a36d.jpg","volume":68,"issue_number":1,"year":2010,"series":null,"season":null,"authors":"Alexander III, John H.","article_content":"A New Plant Introduction from the Arnold Arboretum: Ilex glabra `Peggy's Cove' John H. Alexander III I n October 1988, I was in Nova Scotia for a speaking engagement with the Atlantic Rhododendron and Horticultural Society. Several members were kind enough to show me the sights, including local natural areas. One day we were on a seaside barren, northwest of the fishing village of Peggy's Cove, looking out at the Atlantic Ocean. Crouched between us and the ocean, on a treeless shore that appeared to be more rock than soil, were numerous inkberries, Ilex glabra. I have a special interest in this species and had previously collected inkberry cuttings from the New Jersey pine barrens and from compact plants I spotted while driving along Massachusetts roadways. The plants at Peggy's Cove were dwarfed and misshapen by the harsh seaside environment, growing here near the northern extreme of the species' range. I knew that a plant's response to its environment does not necessarily change its genetic makeup, so cuttings from these dwarfed plants might grow into large, robust shrubs when planted in a favorable garden setting. But then again--they might not. My hope was that, after countless generations growing by the shore, their compact habit was now genetic. As plant propagator's like to say, \"The best time to take a cutting is when it's offered,\" so I collected cuttings from plant after plant until my hosts seemed to grow weary from watching me. Thirty-eight of these cuttings--collected from perhaps 12 different plants--were stuck in the Arboretum's propagation house. At least 19 of the cuttings rooted, and all were given the accession number 929-88. Within this group I found what I had hoped to find: a more compact and smaller-leafed form of Ilex glabra. Named `Peggy's Cove' in honor of its site of origin, this cultivar is not only compact, but it also grows well. It has a mounded habit with branches right to the ground. The latter trait is notable since a complaint sometimes heard about the Ilex glabra cultivars `Densa' and `Compacta' is that they frequently lose their lower branches. Other surviving plants of this collection either didn't grow well or weren't significantly different from cultivars already available. At the Arboretum, the original plant of `Peggy's Cove' (now accession number 500-2007-A) is growing in the Leventritt Shrub and Vine Garden, near specimens of `Compacta' for easy comparison. This 22-year-old plant is now 48 inches (122 centimeters) tall and 60 inches (152 centimeters) wide with a rounded habit. Perhaps it isn't the best example because we have pruned it by harvesting many cuttings from it. Four lightly pruned 8-year-old plants near the Dana Greenhouses (accession number 3-2002) are 22 to 36 inches (56 to 91 centimeters) tall and 30 to 45 inches (76 to 114 centimeters) wide. `Peggy's Cove' is a female, producing the typical small black fruits of this species (if pollinated by a nearby male Ilex glabra). The leaves of `Peggy's Cove' are dark green and smaller than the average inkberry leaf. `Peggy's Cove' inkberry grows best in full sun and tolerates light shade, but may stretch a bit and become less compact if in too much shade. Like many hollies, it prefers acidic soils that are evenly moist. Winter damage to leaves of this evergreen has been slight at the Arboretum. `Peggy's Cove' is probably no hardier than what is typical for the species: USDA zone 5 (average annual minimum temperature -20 to -10F [-28.8 to -23.4C]). It is currently in the early stages of commercial production. Acknowledgments Many thanks to John Weagle, Stanley Dodds, Walter Ostrom, and the late Captain Richard M. Steele. It was these folks who were instrumental in getting me to Nova Scotia and to that barren coast where I found Ilex glabra `Peggy's Cove'. A registration description of this cultivar was published in the Holly Society Journal, 2008, 26(2): 1011. John H. Alexander III is Plant Propagator at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23418","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160af28.jpg","title":"2010-68-1","volume":68,"issue_number":1,"year":2010,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Hill's Oak: The Taxonomy and Dynamics of a Western Great Lakes Endemic","article_sequence":1,"start_page":2,"end_page":14,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25477","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15e856b.jpg","volume":67,"issue_number":4,"year":2010,"series":null,"season":null,"authors":"Hipp, Andrew L.","article_content":"Hill's Oak: The Taxonomy and Dynamics of a Western Great Lakes Endemic KrIS BAchTELL Andrew L. Hipp O aks afford a unique insight into the history of our landscape, flora, and vegetation. Oaks have been among the dominant trees of eastern North American forests and woodlands for approximately 10,000 years (Abrams 1992). Between 8,000 and 3,000 years ago, oaks spread to distributions close to those we observe today (Webb 1981). The landscape between the prairies of the Great Plains and the eastern deciduous forest had by that time settled into a broad transition zone in which prairie, woodland, and savanna shifted with the dynamics of climate and fire (Anderson and Bowles 1999). Some oak species in this region could persist below ground for decades as their shoots were regularly burned to the ground, growing to maturity only when a break in fire frequency allowed their stump sprouts to grow (Kline 1997). The oldest oaks still growing have borne witness to fires, changes in forest structure and composition, and substantial anthropogenic landscape changes. These old oaks sustain The form and fall color of Hill's oak (also known as northern pin oak). large numbers of mammals, birds, and insects. Blue jays, squirrels, and, previously, naturalist to imagine a landscape without oaks. passenger pigeons have eaten, hoarded, and disAt the same time, oaks are remarkable for persed acorns in vast quantities (Johnson and their ability to stump botanists. Even where Adkisson 1986; Keator and Bazell 1998; Price there are only a few species to choose from, we 1999, ch. 1), and civilization rests in part on often struggle to put a name on oaks in the field, the structural and nutritional properties of oaks and annotations on many herbarium specimens (Logan 2005). It is hard for a North American capture decades of disagreement. Oaks are noto- Hill's Oak 3 riously promiscuous, with closely related species able to exchange genes seemingly at will. Pioneering work by James hardin in the 1970s demonstrated hybridization among 14 of the 16 white oak group species of eastern North America, with hybridization occurring almost anywhere that different white oak species grow in sympatry (hardin 1975). In the era of DNAbased taxonomy, hybridization has been demonstrated numerous times using chloroplast and nuclear data (Whittemore and Schaal 1991, Dumolin-Lapegue et al. 1997, curtu et al. 2007, cavender-Bares and Pahlich 2009). For this reason, oaks have been described by two leaders in the field of speciation as a \"worst case scenario for the biological species concept\" (coyne and Orr 2004, p. 43). Our understanding of the depth and orientation of genetic boundaries, our concepts of what constitutes a plant species, and our ability to differentiate morphologically similar species are tangled up in the oaks. A worst case in a genus of worst cases The Western Great Lakes endemic hill's oak (also known as northern pin oak) (Quercus ellipsoidalis; Plate 1) is distinguished by the number of workers who have puzzled over its taxonomic status and proper identification (Trelease 1919; Jensen 1977, 1984; Overlease 1977, 1991; Maycock et al. 1980; Shepard 2009). hill's oak is a member of the black oak group, Quercus section Lobatae, a New World lineage of more than 100 species, of which approximately 75 are found in Mexico and 35 in North America north of Mexico. The section is easily recognized in the field by the presence of bristles or awns on the tips of the lobes (in, for example, Q. velutina, Q. rubra, Q. shumardii, Q. palustris) or leaf apex if the leaf is unlobed (for example, Q. imbricaria, Q. phellos, Q. pumila). Most species in the group also mature acorns over two seasons. In habitat, hill's oak ranges from dry sandstone bluffs, oak barrens, and sand savannas to seasonally wet sandy soils and dry-mesic forests in clayey soils. The tree is particularly common in woodlands of northeastern Illinois. Typical hill's oaks have deeply lobed leaves with moreor-less c-shaped sinuses; leaf undersides that Plate 1. Hill's oak (Quercus ellipsoidalis), showing leaf and acorns. While the smaller leaf size and more ellipsoid acorn are typical of Hill's oak relative to scarlet oak (see Plate 2), leaf and acorn morphology are profoundly variable in Hill's oak. William Trelease (1919) wrote that the \"extremes\" of morphological variation in Hill's oak acorns range continuously from one to the other and have no obvious segregation on the landscape. This is a remarkable statement in light of the fact that the epithet \"ellipsoidalis\" references the acorn shape, which was instrumental in tipping Rev. Hill off to the species' distinctness. Vouchers of the illustrated specimens are deposited at the herbarium of The Morton Arboretum: A.Hipp #3096 (Hoosier Prairie, Lake Co., IN; leaf), A.Hipp & J.Schlismann #2489 (Middlefork Savanna Forest Preserve, Lake Co., IL; acorn). Illustration by Rachel Davis. are smooth or at most sparsely pubescent; terminal buds that are silky-pubescent on the distal (upper) third to two-thirds; and acorn caps that are smooth to sparsely pubescent on the inner surface, with scales on the upper surface that have tightly appressed tips. In these characters, hill's oak is similar to the more widespread eastern North American scarlet oak (Q. coccinea; Plate 2), and in fact it was commonly 4 Arnoldia 67\/4 cially northwestern Indiana, and as a consequence the taxonomy of these two species has remained in flux. We began a study at The Morton Arboretum in 2005 to investigate whether hill's oak, scarlet oak, and the widespread black oak (Quercus velutina; Plate 3) are genetically distinct from ANDrEW hIPP Terminal buds of Hill's oak, showing the silky pubescence on the distal (upper) half of the bud that is typical in this species. Hoosier Prairie, Lake County, Indiana. identified as scarlet oak when first viewed by botanists in the late nineteenth century. In 1891, reverend Ernest J. hill encountered a few populations in the area around Glenwood and calumet Park, cook county, Illinois that he identified as scarlet oak \"with some misgivings.\" With further study, hill judged that the leaf coloration in fall, bark texture, and acorn shape sufficiently distinguished the tree from scarlet oak to warrant its recognition as a separate species, and he published his description of the species in the Botanical Gazette in 1899. Subsequent to this work, many botanists accepted that hill's oak was found throughout the upper Midwest to the exclusion of scarlet oak. however, the distinction between hill's oak and scarlet oak is not always clear. At their morphological extremes, scarlet oak and hill's oak are readily distinguishable. Typical scarlet oak has larger leaves and terminal buds; acorn cap scales with broad, glossy bodies and tips tending to be narrow and somewhat elongate\/ acuminate; and concentric rings of pits around the exposed (stylar) end of the acorn nut that appear as though they were scratched with an etching needle or burned into the acorn. hill's oak has smaller leaves and terminal buds; acorn cap scales with dull or pubescent bodies and relatively short apices; and usually no rings around the tip of the acorn cap, occasionally one or two small rings. But these characters overlap in the greater chicago region, espe- Plate 2. Scarlet oak (Quercus coccinea), showing leaf and acorns; detail of the stylar end of the acorn illustrates the concentric rings typical of this species. While typical scarlet oak does possess these rings, and typical Hill's oak does not, we have found several specimens of Hill's oak that have one ring or, less commonly, two concentric rings of pits at the stylar end of the acorn. In Hill's oak, these rings are mostly solitary when present, 2.753.5 (5) mm in diameter, but in scarlet oak, they are commonly 2 or more and greater than 3.5 mm in diameter. Vouchers of the illustrated specimens are deposited at the herbarium of The Morton Arboretum: A.Hipp & C.Kirschbaum #2627 (Wayne National Forest, Lawrence Co., OH; acorn largely enclosed in cupule, leaf and branch with immature acorns), A.Hipp #3107 (Tinley Creek Forest Preserve, Cook Co. IL; mature acorn, side view and stylar end detail). Illustration by Rachel Davis. Hill's Oak ANDrEW hIPP 5 in northwestern Indiana and southern Michigan that confound our efforts to understand the natural distribution of hill's oak and scarlet oak? Second, do local populations of hill's oak and black oak exhibit gene flow, and does genetic intermediacy between these species correlate with morphological intermediacy? Finally, what is the evolutionary history of black oak section members, and what can this history tell us about the process of oak diversification? Acorns of Hill's oak, illustrating the tightly appressed acorn cap scales that distinguish the species from black oak. Striations on the acorn body are not uncommon in Hill's oak, but also not the rule. Acorn shape in Hill's oak is highly variable. Talltree Arboretum, Porter County, Indiana. one another. My primary collaborator in this project, Jaime Weber, and I have sampled oaks from 58 sites (Figure 1) and genotyped nearly 700 hill's and black oaks as well as populations of scarlet oak from Missouri, southern Illinois, southern Ohio, and upstate New York, and of the related species red oak (Q. rubra), Shumard's oak (Q. shumardii), and pin oak (Q. palustris). We are currently investigating three basic questions. First, are hill's oak and scarlet oak genetically distinct from one another? Do they show the genetic separation we expect of distinct species? can we use genetic data to identify morphologically problematic populations Plate 3. Black oak (Quercus velutina), showing leaf and acorns. The loose apices of the acorn cap scales in typical black oak give the cap a fringed appearance clearly visible in the field. In both Hill's oak and scarlet oak, the acorn cap scale apices are more nearly appressed to the underlying scales, giving the cap a smooth appearance. An important but less recognized character for distinguishing black oak is the pubescence on the inner surface of the acorn cap, which is dense and matted in black oak only (illustrated in Hipp et al. in press). Vouchers of the illustrated specimens are deposited at the herbarium of The Morton Arboretum: A.Hipp #3087 (Hoosier Prairie, Lake Co., IN; leaf), J.Hitz & A.Hipp 100505-13 [TAL-013] (Taltree Arboretum, Porter Co., IN; acorns). Illustration by Rachel Davis. 6 Arnoldia 67\/4 ANDrEW hIPP ANDrEW hIPP Growing in a forest understory, this seedling of scarlet oak (left) shows relatively deep lobing of the leaves compared to those of a black oak seedling (right). Chemung County, New York. Figure 1. Map of species distributions, with sampling localities. The distribution of Hill's oak (Quercus ellipsoidalis) is mapped in dark grey, the distribution of scarlet oak (Q. coccinea) in light grey. Speckling indicates counties in which both species have been reported. Dots indicate sites where species were sampled for the current study. Note that only pin oak (Q. palustris) was sampled from the northern Ohio locality. Base map adapted from Hipp and Weber 2008, with Indiana distribution according to Biagi and Jensen 1995. Hill's Oak 7 Hill's oak and scarlet oak: two different gene pools We began our work uncertain as to whether hill's oak and a genetically distinct scarlet oak were both present in the chicago region. We also did not know whether we would be able to distinguish closely related species at all using genetic data. Previous workers in the region had found that microsatellite data, which is generated by surveying the genome for rapidly evolving repetitive DNA regions, is not consis- tently able to distinguish such species as white oak and its relatives (craft and Ashley 2006) or members of the black oak group (Aldrich et al. 2003). We decided to utilize the amplified fragment length polymorphism (AFLP) technique to genotype trees in this study. The AFLP approach is a shotgun-type approach used for DNA fingerprinting and genome scanning. The method entails cutting the genome of an organism into a large number of pieces at arbitrary points in the genome, then using the size dis- Figure 2. Two-dimensional ordination of 120 individuals representing Quercus coccinea, Q. ellipsoidalis, Q. velutina, and Q. ellipsoidalis x Q. velutina [Q. x palaeolithicola]. The ordination represents the best two-dimensional spatial representation of the genetic distances among individuals. Stated another way, each point on the figure represents a single genotyped oak tree, and the relative proximity between points represents the relative genetic similarity between trees. Ordination methods and voucher numbers are reported in Hipp and Weber 2008. 8 Arnoldia 67\/4 ANDrEW hIPP Scarlet oak trunk, illustrating the planed-off appearance of the bark ridges, reminiscent of (though less pronounced than) red oak. Shawnee National Forest, Gallatin County, Illinois. Hill's Oak 9 ANDrEW hIPP ANDrEW hIPP tribution of the DNA fragments to estimate genetic similarity between organisms. The disadvantages of AFLP data relative to microsatellite and DNA sequence data is that without directly sequencing AFLP markers, one generally has to assume that markers of a given length are identical by descent and that each marker represents a gene region independent of all others sampled, in which we can identify alleles that are present but not alleles that are absent. These facts render the data less useful for population genetic studies than microsatellite data, but the ability Leaf of Hill's oak, illustrating the deep lobing typical of this species and scarto sample large numbers of genes let oak. This specimen (TAG-027, housed at the Herbarium of The Morton across the entire oak genome is Arboretum) genotypes decisively as Hill's oak, but morphologically it appears desirable if we are to detect genetic closer to scarlet oak (see discussion in text of article). Talltree Arboretum, differentiation even in the presence Porter County, Indiana. of interspecific gene flow. All analyses we have conducted demonstrate a strong separation of scarlet oak from the other species investigated, stronger than the separation between hill's oak and black oak (Figure 2). It is important to note that genetic divergence alone does not make a species. It has long been recognized that there can be strong genetic differentiation among populations within species (Ehrlich and raven 1969). however, when genetic divergence between two putative species exceeds genetic differentiation between other closely related taxa recognized as being distinct at the species level, and when this differentiation is associated with geo- Foliage of a putative hybrid between Hill's oak and scarlet oak. This specigraphic distance (allopatry; Figure men (TAG-030, housed at the Herbarium of The Morton Arboretum) is one of the very rare specimens in our study that genotypes as a hybrid between 1), most biologists are inclined to Hill's oak and scarlet oak. These specimens bear further study. Talltree recognize the taxa as distinct spe- Arboretum, Porter County, Indiana. cies. The divergence between scarlet oak and hill's oak must be explained either as there is little association between genetic difdivergence between two species or as genetic ferentiation and geographic distance in black divergence within a single, wide-ranging speoak across a similar geographic range. When cies. Although geographic distance may play a we sample hill's oaks of northwestern Indiana role in the strong separation between these two and southern Michigan that are morphogically species, we have found in follow-up analyses similar to scarlet oak (e.g. Figure 2, individual (hipp and Weber 2008; hipp et al. unpubl.) that TAG-027), for the most part they do not appear 10 Arnoldia 67\/4 to be genetically similar to scarlet oak, though the genotypes of a small number of samples we have collected in northwest Indiana suggest that scarlet oak may be present in that area. It is significant that we find very few individuals with genotypes intermediate between hill's oak and scarlet oak. Naturally-occurring scarlet oak also appears to be rare in the range of hill's oak, with a few exceptions. First, as indicated above, our data suggest that scarlet oak may be present in northwest Indiana, based on a few specimens that are genetically intermediate between hill's oak and scarlet oak. however, the one specimen we sampled from northwest Indiana that appears morphologically to be unambiguous scarlet oak (TAG-027) genotypes as pure hill's oak, and results at other sites where scarlet oak is not present (e.g., central Wisconsin) suggest that occasional genetic assignment discrepancies between hill's oak and scarlet oak may be a consequence of genetic similarity between the two species. Our findings on this bear more detailed follow-up work. Second, we have genotyped a few trees from a stand of scarlet oaks and other southern Illinois trees previously reported from Tinley creek Forest Preserve, cook co., IL (Shepard 2005). Scarlet oaks from this site are the only trees in our study to genotype as pure scarlet oak in the Great Lakes region, with no evidence of introgression from hill's oak or black oak. however, they appear to have been planted in the twentieth century, as they occur on former oldfield habitat (pers. obs.). Moreover, smaller trees from an adjacent ANDrEW hIPP forest margin genotype as scarlet oak as well, though with minimal evidence of introgression from hill's oak, and may be natural offspring of these introduced trees. These facts notwithstanding, the strong genetic disjuncture we see between hill's oak and scarlet oak gives us a great deal of confidence that the morphological intermediacy between them (Shepard 2009) has more to do with intraspecific morphological variation than with gene flow between them. hill's oak and scarlet oak are distinct species. Black oak and Hill's oak: gene flow, but not as much as you might think having determined that hill's oak and scarlet oak are genetically distinct from one another, we were interested in understanding the source of genetic similarity between black oak and hill's oak. In northern Illinois, Wisconsin, and Michigan, distinguishing these two species from each other is not always straightforward. As is the case with hill's oak and scarlet oak, specimens that lie at morphological extremes are easy to identify: typical black oak has large, densely pubescent terminal buds; acorn caps with loose scales and dense, matted pubescence on the inner surface; and leaves that are often pubescent, even roughly so, tending to be less deeply lobed than those of hill's oak. however, morphological intermediates are not uncommon (though with good material they are less common than people may suspect), and our first thought was that morphological intermediacy might be predicted well by genetic intermeANDrEW hIPP Inner surface of a black oak acorn cap (left) shows the matted pubescence typical of the species while the inner surface of a Hill's oak acorn cap (right) is typically hairless. Hill's Oak 11 ANDrEW hIPP ANDrEW hIPP Acorn of black oak, illustrating the loose acorn cap scale tips typical of this species. Talltree Arboretum, Porter County, Indiana. diacy. Our attempt to place morphologically intermediate individuals on our ordinations suggests something different: specimens with mature winter buds and\/or acorns as well as reasonably intact leaves and that nonetheless have characteristics of both hill's oak and black oak genotype across a wide range of the two species rather than in a position intermediate between them (Figure 2). Other researchers have found similar discrepancy between morphological and molecular estimates of admixture (e.g., craft et al. 2002, Gonzalez-rodriguez et al. 2004), which may be a product of the complex history of crosses and back-crosses expected in a group of outcrossing, readily hybridizing species like the oaks. Subsequent analysis of our full set of sampled individuals demonstrates a few misclassifications between black oak and hill's oak, i.e., incongruence between our identifications based on morphology and the population assignments based on genetic data: 14 black oak out of 286 sampled have > 0.20 assignment to hill's oak in a commonly used Bayesian population genetic analysis approach. This mismatch between genetic and morphological species assignments is a hallmark of introgressive hybridization and has been reported previously in oaks (cavenderBares and Pahlich 2009), and the presence of such individuals supports the hypothesis of gene flow between the two species. It is remarkable, however, that we find so little genuine misclassification or evidence of genetic admix- Branch of black oak, illustrating the densely pubescent buds typical of the species. Black oak has distinctive yellow petioles at some sites, as illustrated here, but that character is not reliable in much of the range of the species (though in The Trees of Vermont by Burns and Otis (1916), petioles of black oak are described as \"stout, yellow, 3 to 6 inches long\"). Talltree Arboretum, Porter County, Indiana. ture between black oak and hill's oak. Our findings build on those of a now-classic study of European oaks (Muir et al. 2000) in demonstrating that while oaks do hybridize, there are enough barriers to interspecific gene flow to make oak taxonomy a meaningful enterprise. Phylogeny of the black oaks: a little information, a lot to learn Our work going forward is aimed at understanding how these species and their relatives are related, and how contemporary gene flow and evolutionary history interact to define the limits of today's oak species. Utilizing a larger AFLP dataset and species sampling, we have found that hill's oak and scarlet oak are sister 12 Arnoldia 67\/4 ANDrEW hIPP View from High Knob, overlooking a forest of white and scarlet oak. Shawnee National Forest, Gallatin County, Illinois. Hill's Oak 13 species, meaning that they share a more recent common ancestor than either shares with black oak, red oak, pin oak, or any other species. The morphological overlap we see between hill's oak and scarlet oak suggests that the two species may have inherited a similar pool of characteristics from a recent common ancestor, though these characteristics were inherited in differing proportions. This finding is particularly interesting in light of the distribution of hill's oak and scarlet oak. hill's oak is the only oak species endemic to the Great Lakes region (Abrams 1992) and is distributed almost exclusively in glaciated terrain. It is tolerant of disturbance and has been characterized as the most drought-tolerant of the black oak species (colodonato 1993), though it appears to be less common than black oak in the driest sand soils of northern Illinois. Its geographic range also overlaps closely with the distribution of dry soil oak savannas in the Great Lakes region (Will-Wolf and Stearns 1999). Scarlet oak, on the other hand, is distributed predominantly south of the edge of the ice sheet at the last glacial maximum. While also tolerant of disturbance and favoring dry sandy or gravelly soils, scarlet oak is not uncommon in mature forests in more mesic soils (carey 1992). Given the broad geographic extent of scarlet oak and the compressed distribution of hardwood forests during the glacial maximum (Delcourt and Delcourt 1984), these two species likely co-occurred for at least a portion of the Pleistocene. Why, then, has hill's oak migrated into postglacial environments while scarlet oak is largely confined to unglaciated terrain? It may be that differences in cold tolerance between the two species govern their relative distributions. hill's oak may also be more tolerant of disturbance or of higher ph or finer soil texture. If so, it may have been more able to take advantage of newly opened territory as the vegetation of the savanna regions around the Great Lakes shuffled around rapidly following glacial retreat. This capacity to respond to relatively rapid environmental change may bode well for hill's oak in the future. In the shorter term, our growing understanding of oak evolutionary relationships and ecology should allow us to address basic questions about oak distribution and speciation, and guide predictions about how tree species will respond to future climatic and environmental changes. Acknowledgements This work has been funded predominantly by The Morton Arboretum, where oaks have been a priority in research and the living collections for decades. Additional funding was provided by grants from The American Philosophical Society and The hanes Fund of the Michigan Botanical club. rachel Davis, Marlin Bowles, and Paul Manos generously provided feedback on a draft of this article. References Abrams, M.D. 1992. Fire and the development of oak forests. BioScience 42: 346353. Aldrich, P.r., G.r. Parker, c.h. Michler, and J. romeroSeverson. 2003. Whole-tree silvic identifications and the microsatellite genetic structure of a red oak species complex in an Indiana old-growth forest. Canadian Journal of Forest Research 33: 22282237. Anderson, r.c., and M.L. Bowles. 1999. Deep-soil savannas and barrens of the Midwestern United States. In: r. c. Anderson, J. S. Fralish and J. M. Baskin, eds. Savannas, barrens, and rock outcrop plant communities of North America, pp. 155170. cambridge University Press, cambridge. Biagi, A. and r.J. Jensen. 1995. The genus Quercus (Fagaceae) in Indiana: Phytogeography and a key to the species. Indiana Academy of Science 104: 1124. carey, J.h. 1992. Quercus coccinea. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, rocky Mountain research Station, Fire Sciences Laboratory (Producer). Available: http:\/\/www.fs.fed.us\/ database\/feis\/plants\/tree\/quecoc\/all.html [Accessed 5 January 2010]. cavender-Bares, J. and A. Pahlich. 2009. Molecular, morphological, and ecological niche differentiation of sympatric sister oak species, Quercus virginiana and Q. geminata (Fagaceae). American Journal of Botany 96: 16901702. coladonato, M. 1993. Quercus ellipsoidalis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, rocky Mountain research Station, Fire Sciences Laboratory (Producer). Available: http:\/\/www. fs.fed.us\/database\/feis\/plants\/tree\/queell\/ all.html [accessed 5 Jan 2010]. craft, K.J., M.V. Ashley, and W.D. Koenig. 2002. Limited hybridization between Quercus lobata and 14 Arnoldia 67\/4 Quercus douglasii (Fagaceae) in a mixed stand in central coastal california. American Journal of Botany 89: 17921798. craft, K.J. and M.V. Ashley. 2006. Population differentiation among three species of white oak in northeastern Illinois. Canadian Journal of Forest Research 26: 206215. curtu, A., O. Gailing, and r. Finkeldey. 2007. Evidence for hybridization and introgression within a species-rich oak (Quercus spp.) community. BMC Evolutionary Biology 7: 218. Delcourt, h. r., and P. A. Delcourt. 1984. Ice Age haven for hardwoods. Natural History 93: 2228. Dumolin-Lapegue, S., et al. 1997. 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The affinities and misbehaviors o f h i l l 's o a k ( Q u e r c u s e l l i p s o i d a l i s ) . International Oak Journal. Jaccard, P. 1908. Nouvelles recherches sur la distribution florale. Bulletin de la Societe Vaudoise des Sciences Naturelles 44: 223270. Jensen, r.J. 1977. A preliminary numerical analysis of the red oak complex in Michigan and Wisconsin USA. Taxon 26: 399407. Jensen, r.J., r. Depiero, and B. K. Smith. 1984. Vegetative characters, population variation, and the hybrid origin of Quercus ellipsoidalis. American Midland Naturalist 111: 364370. Johnson, W.c. and c.S. Adkisson. 1986. Airlifting the oaks. Natural History 10\/86: 4146. Keator, G. and S. Bazell. 1998. The life of an oak: An intimate portrait. heyday Books, Berkeley. Kline, V. M . 1997. Orchards of oak and a sea of grass. In: S. Packard and c. F. Mutel, eds. The Tallgrass restoration handbook for prairies, savannas, and woodlands, pp. 321. Island Press, Washington, D.c. Logan, W. B. 2005. Oak: Frame of civilization. W.W. Norton & company, Inc., New York. Maycock, P. F., D. r. Daniel, r. Gregory, and A. A. reznicek. 1980. hill's oak (Quercus ellipsoidalis) in canada. Canadian FieldNaturalist 94: 277285. Muir, G., c.c. Fleming, and c. Schlotterer. 2000. Species status of hybridizing oaks. Nature 405: 1016. Overlease, W. r. 1977. A study of the relationship between Scarlet oak (Quercus coccinea Muenchh.) and hill oak (Quercus ellipsoidalis E.J. hill) in Michigan and nearby states. Journal of the Pennsylvania Academy of Science 51: 4750. Overlease, W.r. 1991. Genetic relationships between three species of oaks as deter mined by common garden studies with populations from Michigan, Indiana and Wisconsin. Journal of the Pennsylvania Academy of Science 65: 7174. Price, J. 1999. Flight maps: Adventures with nature in modern America. Basic Books, Perseus Books Group, New York. Shepard, D.A. 2005. The land that time forgot: Southern flatwood oaks and associates of the Tinley creek Forest Preserve of cook county, Illinois. International Oak Journal 16: 4760. Shepard, D.A. 2009. A review of the taxonomic status of Quercus ellipsoidalis and Quercus coccinea in the Eastern United States. International Oak Journal 20: 6584. Trelease, W. 1919. The jack oak (Quercus ellipsoidalis). Transactions of the Illinois State Academy of Science 12: 108118. Webb, T. 1981. The past 11,000 years of vegetational change in eastern North America. BioScience 31: 501506. Whittemore, A.T. and B.A. Schaal. 1991. Interspecific gene flow in sympatric oaks. Proceedings of the National Academy of Sciences of the United States of America 88: 25402544. Will-Wolf, S. and F. Stearns. 1999. Dry soil savanna in the Great Lakes region. In: r. c. Anderson, J. S. Fralish and J. M. Baskin, eds. Savannas, barrens, and rock outcrop plant communities of North America, pp. 135154. cambridge University Press, cambridge. Andrew L. hipp is Plant Systematist and herbarium curator at The Morton Arboretum in Lisle, Illinois. rachel Davis is an artist in Downers Grove, Illinois. "},{"has_event_date":0,"type":"arnoldia","title":"Tree Hormones and Why They Matter","article_sequence":2,"start_page":15,"end_page":19,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25479","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed060a328.jpg","volume":67,"issue_number":4,"year":2010,"series":null,"season":null,"authors":"Murray, Joseph","article_content":"Tree Hormones and Why They Matter Joseph Murray T rees are the oldest, largest, and perhaps the most complex organisms on earth. Increasingly, society has moved beyond simply appreciating trees for the beauty and shade they offer, and now recognizes the significant societal, environmental, and economic benefits trees provide. These benefits can be especially important in urban areas, yet many urban sites present very difficult situations for growing trees. Most tree species should be able to live and provide benefits for several hundred years, but urban trees--often plagued by poor soil, restricted root zones, and limited care--rarely achieve even a fraction of their potential life spans. The more we (arborists, city foresters, growers, etc.) know about the biology of trees, the better we will be able to apply proper arboricultural practices to help trees help themselves. Plant hormones and their effect on tree behavior is an often overlooked aspect of arboriculture. Plant hormones--generally defined as sub- The hormone pathway runs from roots to branch tips in trees such as this stances produced in very small amounts Stewartia pseudocamellia. in the plant that influence the plant's concentration, or interaction--for everything physiological processes--play a crucial role in happening in trees. Today, most plant science helping the plant to make adjustments in a textbooks describe five major plant hormones: changing environment. Knowing more about auxin, cytokinin, gibberellins, abscisic acid, and how plant hormones work in trees helps in ethylene. However, there are more than five understanding the implications of such comhormones in plants and research is ongoing. mon arboricultural practices as pruning, plantPlant hormones present a number of chaling, fertilization, and irrigation. lenges to the physiologists attempting to What Do We Know About Plant Hormones? understand how they operate. Plant hormones Prior to 1950 in the United States, this artiare produced, and are active, in very small cle would only have addressed two hormones, concentrations. At different times during the auxin and ethylene, which were then considgrowing season, different parts of the plant ered responsible--by their presence, absence, produce specific hormones that influence dis- PeTer Del TreDIcI 16 Arnoldia 67\/4 NANcy roSe Hormones and Tropisms An interesting example of a hormone causing a plant response is auxin's role in phototropism. A tropism in plants is any growth response resulting in the curvature of a plant organ toward or away from stimuli. Phototropism in plants typically consists of new growth in the shoot system growing toward light. Light striking the side of new growth at the end of branches stimulates the tissues to produce auxin, which then migrates to the opposite (dark) side of the stem where it triggers a physiological response loosening longitudinal cell walls, allowing those cells to expand in length, thus resulting in the curvature of the stem toward the light. Similarly, gravitropism also results in the curvature of the new growth in the root system downward to gravity in response to the unequal cell expansion in the tissue just behind the root tip. In addition to light and gravity, there are many other forms of stimuli that elicit a growth response. This Hippeastrum exhibits phototropism--plant growth bending toward light. Charles Darwin was one of the first to research the mechanics of phototropism and, with his son Francis, published a summary of their observations in the book The Power of Movement in Plants in 1880. Later researchers identified auxin as the plant hormone involved in phototropism. tant tissues that are receptive for brief periods of time. Furthermore, the same hormone may cause two different responses in the same receptive tissues, depending upon the concentration of the hormone. Hormones are signal transducers, converting an environmental stimulus into a physiological or anatomical response. As an example, let's look at how sunlight makes roots grow in the spring, via a simple pathway using the plant hormone auxin. It makes sense for a tree to invest resources into the root system before the shoot system, so early in the spring sunlight on the shoot apical meristem (bud) and young leaves results in these tissues producing auxin, which travels down to the roots. Hormones in plants may travel throughout the plant but will only affect tissues composed of cells that have special receptors to receive that particular hormone. These target cells may perform a number of functions in response to the arrival of the plant hormone. In a physiological response similar to that described for phototropism (see textbox), auxin stimulates cells at the root tips to release hydrogen ions into the surrounding cell walls. In response to the decreasing pH, enzymes become activated and begin loosening bonds between cellulose microfibrils, thus softening the cell walls. Inside the plant cell is an organelle, the central vacuole, full of water that is continually pressing against the cell wall resulting in turgor pressure. The collective action of softened cell walls expanding in response to the central vacuoles results in the elongation of the root tips. The signal transduction is complete. The hormone auxin allowed the tree to translate an environmental stimulus into a physiological and anatomical response. Simply put, sunlight made roots grow. The Auxin-Cytokinin Pathway Many gardeners are familiar with a common technique to produce bushier plants; by simply pinching off the end of a growing stem, Hormones 17 NANcy roSe responsible for the auxin's production. left out of this pathway are the numerous lateral buds, especially those near the end of the branch. Without receiving the spring wake-up call from cytokinin, these lateral buds become dormant. Although they are no longer visible at the surface, each year the dormant buds move outward with the vascular cambium so that they remain close to the surface. Should something happen to disrupt the auxin-cytokinin pathway, then they may emerge and grow into branches, setting up their own auxin-cytokinin pathways with the root system. It's also important to recognize that there are specific enzymes located at the shoot and root tips to destroy the arriving hormones after they have had their effect. These hormone-destroying enzymes are produced in the same tissue near the shoot and root tips. In the root tips, an enzyme is produced that will destroy auxin, just as in the shoot tips, an enzyme is made to destroy cytokinin. Should these enzymes not perform their tasks, the concentration of hormones will increase and cause a different response in the receptive tissues. Removal of the branch tip (center of photo) disrupted the auxin-cytokinin pathway, allowing lateral shoots to develop just below the removal point. there is a proliferation of branch development below the area that was removed. This growth response demonstrates what happens when the auxin-cytokinin pathway is disrupted. The downward flow of auxin creates a pathway from the terminal buds to the root tips. As mentioned, the auxin acts as a signal transducer, notifying the roots that it's spring and it would be in the best interest of the tree to begin growing roots for the season. In addition to growth, the tissues in the root tips produce the hormone cytokinin. cytokinin, like auxin, is going to stimulate growth as well, but in a different location--at the ends of the very branches that originally established the auxin pathway. each spring, the auxin-cytokinin pathway promotes the timely growth of the root and shoot systems. like a male insect following a pheromone trail produced by a receptive female insect, cytokinin follows the increasingly stronger gradient of auxin directly to the shoot tips Common Tree Care Practices and the Impact of Hormone Pathways Knowing that plants have internal mechanisms helping them with an ever-changing environment should make us pause and attempt to understand what is happening in the plant before beginning to actively \"care\" for the plant. Sometimes our efforts at achieving short term goals (e.g., darker green foliage, more growth, controlled shape) may be aggravating the tree's ability to achieve optimal health. Trees' hormone pathways are involved in the arboricultural practices described below: Transplanting regardless of how carefully balled-andburlapped or container-grown trees are transported and installed, some roots will be damaged and die. The roots that are particularly susceptible to damage are the very fine root tips. And it is these same roots that are to produce cytokinin and transport it up to the shoot tips to stimulate elongation of branches. This is why newly transplanted trees are so slow at developing significant shoot growth during 18 Arnoldia 67\/4 NANcy roSe be the dominant hormone directing the majority of resources to continue root growth, and a larger root system enables a search through a greater soil volume for nutrients. In nutrient poor soil, it is in the tree's best interest to invest its limited resources in root growth and not shoot growth. But if a fertilizer is applied, the root system is fooled into thinking it is in a nutrient-rich environment and the production of cytokinin increases, resulting in a larger shoot system relative to the root system. If this nutrient subsidy ceases, the tree is caught with a shoot system that cannot be sustained with the current root system. Irrigation cytokinin also functions in the opening of stomata on the underside of leaves, allowing the steady movement of water from the roots to the leaves. The arch-rival of cytokinin is another root-derived hormone called abscisic acid. Abscisic acid is responsible for the closure coUrTeSy oF BoB MUgAAS Auxin accumulates at the base of stem cuttings, stimulating root initiation. Exogenous auxins, in the form of rooting powders or dips, are often applied to the bases of woody plant stem cuttings before sticking in propagation beds (rooted Microbiota decussata cuttings seen here). the first year or two after transplanting. The loss of the root tips also means the loss of the ability to produce the auxin-destroying enzymes. As a result, the auxin concentration increases until the surrounding tissue responds by generating adventitious root growth. This kind of root proliferation can be observed when an African violet leaf stem is placed in water. Auxin moves down the base of the stem until it builds in concentration at the point the stem was severed from the plant, changing stem tissue into actively growing root tissue. Fertilization So long as there is adequate nitrogen available in the soil, tree roots will continue producing cytokinin at the appropriate times of the year in response to the establishment of the auxin pathway. However, when the nitrogen level is inadequate, the root system will suspend cytokinin production. Auxin will then When trees receive environmental subsidies, such as supplemental water from lawn irrigation systems, their internal regulatory mechanisms can be disrupted resulting in imbalanced root-to-shoot growth. Hormones 19 JoSePH MUrrAy pathway. Should the shoot tips be removed, the timely production of auxin and its transport to the roots will not occur in the spring. This means the cytokinin produced in the roots will not know where to travel to stimulate the growth at the end of the branch. The concentration of cytokinin will increase at the point where the branch broke or was cut because the tissue responsible for producing the cytokinin-destroying enzymes is gone. As a result, cytokinin will spread through this new truncated terminal end of the branch, finding and releasing the latent buds. This is why there is a proliferation of watersprouts emerging at the end of branches damaged by storms or by the ill-advised practice of topping trees. Topping, an improper pruning practice in which tree trunks and major branches are drastically cut back, results in a proliferation of weakly attached lateral shoots at the pruning points. of the stomata when there is not enough soil moisture to perform photosynthesis. As long as the fine roots are in contact with soil and able to absorb water, cytokinin is being produced and traveling to the leaves to keep the stomata open. Should the soil begin to dry and soil particles pull away from the roots, the root system will produce abscisic acid and send it to the foliage to shut the stomata. Periodic episodes of landscape irrigation disrupt this internal regulatory mechanism, possibly placing those irrigated trees at risk for more severe damage. If periodic irrigation stops (perhaps from failure of an irrigation system or institution of municipal watering bans) the trees are suddenly exposed to drought conditions made even more acute because the shoot system has developed at a faster pace than the root system. Lessons Learned Trees have existed for over 300 million years. The evolution of a hormone system allowed early plants to deal with a changing environment and to coordinate their parts in time and space. And for venerable trees, these hormone systems are particularly important. As caretakers of trees in urban areas, it is our duty to first understand these subtle internal mechanisms before blithely applying a treatment that we believe is in the tree's best interest. Bibliography Davies, P.J. 1995. The Plant Hormones: Their Nature, occurrence, and Functions. In: Plant Hormones: Physiology, Biochemistry and Molecular Biology. Kluer Academic Publishers. Boston. Kozlowski, T.T. 1979. Tree Growth and Environmental Stresses. University of Washington Press, Seattle. raven, P.H., g.B. Johnson, J.B. losos, and S.r. Singer. 2005. Biology. 7th edition. Mcgraw Hill. Boston Taiz, l. and e. Zeiger. 2006. Plant Physiology. 4th edition. Sinauer Associates, Inc., Sunderland, Massachusetts Wareing, P.F. 1974. Plant Hormones and crop growth. Journal of the Royal Society of Arts. 122: 818827. Joseph Murray is an Assistant Professor of Biology at Blue ridge community college, in Weyers cave, Virginia. He is an International Society of Arboriculture certified arborist, certified utility arborist, and a certified tree worker\/climber specialist. Improper Pruning Cuts or Storm Damage Similar to the response observed in trees following transplanting, the loss of shoot (branch) tips will also disrupt the auxin-cytokinin "},{"has_event_date":0,"type":"arnoldia","title":"2009 Weather at the Arboretum","article_sequence":3,"start_page":20,"end_page":24,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25475","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15e816d.jpg","volume":67,"issue_number":4,"year":2010,"series":null,"season":null,"authors":"Famiglietti, Bob","article_content":"2009 Weather at the Arboretum Bob Famiglietti A s in 2008, greater than normal rainfall occurred in 2009, resulting in optimum soil moisture conditions at the Arboretum. Excellent growth rates were recorded on a vast majority of our woody plants. January was colder than normal. The minimum temperature dropped to at least 28F on every night, a rare occurrence. Readings of -1F were recorded on the 15th, 16th, and 17th, the low for the year. Three storms that each deposited about 6 inches of snow left a persistent snow cover on the ground. Snow total for the month was 21 inches. February was mild and dry with only 7 inches of snow, six of that coming on the 3rd. The relative warmth reduced a foot of accumulated snow on the ground at the beginning of the month to all but a trace by the end of the month. February's high temperature of 60F was reached on the 27th. March had average temperatures and produced only 10 inches of snow. A temperature of 61F occurred twice, and the snow pack melted by mid month. april was a month of extremes. It began cool, as low temperatures dropped into the 30s for thirteen days. Our last freeze occurred on the 13th when it hit 32F. Temperatures soared to the other extreme by the end of the month. Our first day over the 70F mark arrived on the 24th, making it to 71F. It reached 86F on the 25th and 26th and then soared to 95F on the 28th, an amazing leap from the freezing temperature barely two weeks earlier. This was the highest temperature since June 2008, and also turned out to be the high for the year. Rainfall was 4.13 inches for the month. May was warm, cloudy, and dry. Even though rain was measured on fourteen days, it only totaled 2.76 inches for the month. Weather conditions for the Arboretum's annual Lilac Sunday event on May 10th were extremely windy, with gusts over 40 miles per hour. A high of 91F was reached on the 21st, the only reading in the 90s for May. June had eighteen consecutive days with below normal temperatures (8th25th) finishing almost 5F below normal for the month. It was the third coldest June in 183 years of Boston weather-keeping records. Clouds were persistent and rain was measured on nineteen days with traces on four others. Precipitation was 3.99 inches for the month and there were only six days when no water was detected in our rain gauge. A frequent east wind kept us cloudy, cool, and damp. These cool, damp, early summer conditions made it an excellent year for post-transplanting establishment of new plants in the collection; little supplemental watering was needed. On the negative side, the cool, damp weather exacerbated a widespread outbreak of the late blight fungus (Phytophthora infestans) in the Northeast. Late blight attacks plants in the nightshade family (Solanaceae) and is the fungus that was a major factor in the Irish potato famine of the 1850s. Farmers and home gardeners in the region had to destroy tomato and potato crops to prevent the spread Weather 21 Arnold Arboretum Weather Station Data 2009 avg. Max. (F) Jan Feb Mar apr May Jun Jul aug Sep OcT nOv Dec 29.8 39.6 44.1 60.1 68.7 71.5 78.0 82.2 71.1 58.2 55.4 39.4 avg. Min. (F) 13.9 21.6 27.7 40.1 50.3 55.5 61.6 65.2 52.8 41.2 40.7 23.7 avg. Temp. (F) 21.8 30.6 35.9 50.1 59.5 63.5 69.8 73.7 62.0 49.7 48.1 31.6 Max. Temp. (F) 40 60 61 95 91 83 88 93 79 73 69 69 Min. Temp. (F) -1 3 8 30 43 43 51 55 41 32 29 9 precipi- Snowtation fall (inches) (inches) 4.65 2.07 3.01 4.13 2.76 3.99 7.91 3.40 3.28 5.62 3.76 5.27 10.5 21.0 7.0 10.5 average Maximum Temperature . . . . . . . . . . 58.2 average Minimum Temperature . . . . . . . . . . 41.2 average Temperature . . . . . . . . . . . . . . . . . . . 49.7 Total precipitation . . . . . . . . . . . . . . . . . . . . . 49.85 inches Total Snowfall. . . . . . . . . . . . . . . . . . . . . . . . . 49.0 inches Warmest Temperature . . . . . . . . . . . . . . . . . . 95 on april 28 coldest Temperature . . . . . . . . . . . . . . . . . . . -1 on January 15,16, and 17 last Frost Date . . . . . . . . . . . . . . . . . . . . . . . . 32 on april 13 First Frost Date . . . . . . . . . . . . . . . . . . . . . . . . 32 on October 19 growing Season . . . . . . . . . . . . . . . . . . . . . . . 189 days 22 Arnoldia 67\/4 SuE A. PFEIFFER a lightning strike at about 9 a.m. on July 2, 2009, destroyed this venerable nikko fir (Abies homolepis) in the arboretum's conifer collection. The explosive force threw pieces of the tree at least 180 feet away. of late blight. The Arboretum has very limited holdings of woody plants in this family and no collections plants were affected. The damp conditions were also a factor in the appearance of fire blight (Erwinia amylovora), a bacterial disease, on some rose family (Rosaceae) plants in the collections. A high temperature of 83F (lower than in April or May) was reached on the 26th. July was also cloudy, cool, and wet, with 7.91 inches of rain, the sixth wettest July on record. There were fourteen days with measurable rainfall and traces on four others. Thunderstorms were frequent; on the 2nd, a lightning strike during a thunderstorm destroyed a notable 91-foot-tall, 110-year-old Nikko fir (Abies homolepis) in the Arboretum's conifer collection. 2.93 inches of rain fell on the 23rd, the highest one day total since December 11th, 2008. For five days it remained in the 60s and on eleven days it never made it out of the 70s. A high of 88F was recorded on the 18th and 28th. We never reached 90F, which is extremely rare for July. The combined JuneJuly average temperature was the 4th coldest in Boston's recorded weather history. auguST was very warm and, with only 3.4 inches of rain, our driest summer month. Measurable precipitation was recorded on only eight days. The high of 93F was reached on the 18th. 90F or greater was recorded on the 17th through the 19th, creating our only official heat wave of the summer. Weather NANCy ROSE 23 visitors and arboretum staff commented on the outstanding orange-russet fall color exhibited by the dawn redwoods (Metasequoia glyptostroboides) near the hunnewell visitor center late in the autumn of 2009. 24 Arnoldia 67\/4 SepTeMber was cool, sunny, and a bit dry. A heavy rain occurred on the 11th and 12th, but rainfall was measured on only five days for a total of 3.28 inches. Long sunny breaks occurred between rain days. No temperatures of 80F or higher were recorded during the month. OcTOber was cold and wet. Our growing season ended on the 19th with a low of 32F. This was the 21st coldest October in 138 years of Boston weather history. Precipitation was measured on fifteen days for a total of 5.62 inches. Damp conditions notwithstanding, visitors to the Arboretum enjoyed another great fall foliage display this year. KEvIN B. SChOFIELD a cool October followed by unusual warmth in november triggered an abundance of premature late-autumn blooms on this Fuji cherry (Prunus incisa f. serrata) in the bradley rosaceous collection. an early December snow brought an end to the spring preview. nOveMber was warm and somewhat dry, ranking as the 7th warmest November on record. It was only slightly cooler than October. A high of 69F was recorded on the 9th. A low of 29F was recorded on the 6th and 17th. This warm weather kept containerized nursery plants at the Arboretum's Dana Greenhouse from going completely dormant, the condition needed for winter root cellar storage. Many of our containerized plants had to wait for the cold of December to drop their leaves. Though they commonly open a few blossoms during late fall warm-ups, this year some of the mature cherry (Prunus spp.) trees in the Bradley Rosaceous Collection appeared to be in nearly full bloom. DeceMber started warm, reaching a high of 69F on the 3rd. But it then turned cold, remaining below freezing for eight straight days from the16th through the 23rd. This is just what our containerized woody plants needed to go into dormancy, and they could finally be put to bed for the winter. Almost a foot of snow fell over the weekend of the 19th and 20th. Bob Famiglietti is a horticultural Technologist at the Arnold Arboretum's Dana Greenhouses. "},{"has_event_date":0,"type":"arnoldia","title":"An Essay on Naming Nature: The Clash Between Instinct and Science","article_sequence":4,"start_page":25,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25476","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15e8526.jpg","volume":67,"issue_number":4,"year":2010,"series":null,"season":null,"authors":"Stevens, Peter","article_content":"An Essay on Naming Nature: The Clash Between Instinct and Science P. F. Stevens Naming Nature: The Clash Between Instinct and Science Carol Kaesuk Yoon. W.W. Norton & Company, 2009. 344 pages. SBN 978-0-393-06197-0 S ystematics, the science of the study of relationships between organisms, has seen remarkable developments over the last fifty years. Carol Kaesuk Yoon was a graduate student in Cornell in the late 1980s, trying to elucidate the relationships between some fruit flies using the then still fairly novel technique of DNA analysis. There she witnessed some of the vitriolic debates between cladists and evolutionary biologists, two warring groups of systematists who interpreted relationships in very different ways. In fact, analysis of the molecular data that she and others were then starting to use has had profound consequences for our understanding of the living world, and our knowledge of the genealogical relationships between organisms is increasing by leaps and bounds. Taxonomists, those who classify, have in many cases redrawn the limits of groups to better reflect these genealogical relationships. Of course, systematists had long been interested in such relationships, but they used morphological differences to establish them. As Ernst Mayr (who figures in the book's pages) noted, everybody could tell a toucan, with its remarkable beak, from a barbet. Brightly colored though the latter bird might be, barbets had much more conventional bills, and nobody in his or her right mind would put toucans and barbets in the same family. But that is exactly what the genealogical evidence suggested to some. The resolution of this particular story is that barbets are now in four separate families, toucans remaining in their own family. For some, this is a satisfactory solution; after all, this taxonomy does take into account genealogy. But situations like these seem to make no sense intuitively--are birds to be included in reptiles, are we humans really to be placed with fish, as genealogy would suggest? Such questions led Yoon to reflect on where taxonomists and systematists were going. They seemed to have taken leave of their everyday senses as they peered myopically at bands on gels that represented DNA. On the other hand, we have always classified the living world using our 26 Arnoldia 67\/4 ordinary senses, and these classifications make that world real to us in a way that the new classifications do not. It is this world--she calls it the umwelt, the world as it is apparent to our senses, the natural order that it discloses to us--that matters to us. In the world as we perceive it, objectivity, hypothesis testing, and evolutionary change are not relevant; the whale is a fish of sorts, as are clams and maybe even coots, and humans are not apes. This is folk taxonomy, not a scholarly endeavor but a hard-wired and ageless tradition that was coopted by Linnaeus and hijacked by molecular systematists. In the book, we then embark on a fascinating tour. Linnaeus's Herculean labors in classifying the world are explained in detail, \"capturing,\" as he did, \"the essential vision of the living world . . . the vision of the human umwelt\" (p. 50). A brief discussion on Darwin's barnacles ends with the conclusion that all his brilliant evolutionary inspirations would cripple taxonomy--a wall was being erected between the scientist and the living world. Indeed, despite the title of his book, On the Origin of Species . . . , arguments about what a species really was were not settled by Darwin, nor later by Mayr, who thought that because he and New Guinea tribesmen could recognize the same species of birds this made species objectively real. This observation simply made James Watson wonder why Harvard faculty were needed to name things if they did no better than New Guineans. The classifications of plants and animals all over the world show remarkable cross-cultural similarities, down to the numbers of different things that are included in any one classification--which turns out to be similar to the number of genera that some of Yoon's informants, professional taxonomists, could remember; around 600 is the upper limit. Similar numerical regularities apply to species; few genera have more than seven species. Indeed, there are general memory rules here, as George Miller noted in his classical paper, \"The Magical Number Seven, Plus or Minus Two: Some Limits on Our Capacity for Processing Information.\" The great taxonomist George Bentham was aware of such limits as he wrote Genera plantarum with Joseph Dalton Hooker in the later nineteenth century. Yoon notes that it has been shown that names of fish sound like names of, well, fish, rather than of birds or some other animal. This is the classificatory umwelt that we have left behind. She also describes some remarkable people with brains damaged in particular ways who could no longer classify organisms. Returning to academia, the arguments between the three main group of systematists--cladists, pheneticists, and evolutionary systematists--are described very perceptively. Yoon sees that the cladistic approach--recognizing relationships because of shared unique characters--has allowed us to start assembling a tree of life that shows us surprising things about the world. However, this is not the world of our senses, since the living world has been excluded. What is the mere mortal to do? Indeed, there is a tension here. Yoon suggests that classifications were developed specifically for communicating about organisms. However, classifications extend to every part of our world, living or not. We classify items in a supermarket just as effectively as we do organisms. We may have lost contact with life, but we have not lost the ability to classify. Indeed, classification is not so much part of an umwelt that has to do with life in particular but something we do to everything. The binomial, a noun-adjective combination that Linnaeus used, is simply two words we use to describe groups of things, whatever they may be. A red cart and a red oak have the same grammatical and cognitive structure, but one refers to things and the other to plants. In the end, Yoon suggests that we name organisms as we please. There is no one classification, but each classification is a variation on a universal theme; we must reclaim our own umwelts. And herein is food for thought. What is our umwelt? She acknowledges that all individual classificatory systems may be different, but of course the great advance made almost inadvertently by Linnaeus was a way of communicating. A common language, a common classification, is always essential. And whether our umwelt tells us anything in particular or stable is debatable, certainly, our attitudes to the environment have changed dramatically over the last few hundreds of years, and our prelapsarian ideas might not seem very satisfactory to us now. Eyewitness accounts may well be decidedly less than accurate, as any trial Naming Nature 27 NANCY ROSE Carolus Linnaeus, a larger-than-life bronze statue of the \"father of modern taxonomy\" by Robert Berks, in the Heritage Garden at the Chicago Botanic Garden. lawyer or judge will know. Thus, to oppose the new classifications we are developing with an umwelt-based classification that reflects an understanding of the phenomenal (= real) world, seems a mistake. I have been through the biological battles that Yoon describes, and am also a maker, user, and teacher of classifications. There is much more than just DNA sequencing and changing names going on. We are learning much more about the living world and in such a way that it makes us wonder and understand in a way that was impossible before. When I take beginning biology students around the campus and talk about bacteria in the nodules in the pea family, and the bacteria-that-were that pervade cells as mitochondria and chloroplasts (all features that also reflect the new classifications) students clearly understand the world in a very different way. A classification based on umwelt and instinct would be a sorry substitute. The reader will learn a great deal from this book, which is well and clearly written (although the asparagus has never been included in the orchid family, p. 235). The issues that it raises are ongoing. Even aside from the \"debate\" over global warming and evolution, scientists sometimes forget the limits of their world: their truth is not necessarily broadly self-evident. Readers of Yoon's book will surely enjoy Trying Leviathan: . . . by D. Graham Burnett, which raises similar issues, but in a historical context, as the subtitle of that book explains: The Nineteenth-Century New York Court Case That Put the Whale on Trial and Challenged the Order of Nature. The ultimate question is surely why we need alternative classifications and what are the situations in which they help--and what are those in which they are a hindrance. Whether the umwelt, whatever it is (and the word is overused in this review as it is in the book itself), will help us as we think about this, I do not know, although I doubt it. And we do need to think about what is, not what seems--and I say this fully aware of the difficulties surrounding that most simple of words, \"is\". P. F. Stevens is professor of biology at the University of Missouri, St. Louis, and curator, Missouri Botanical Garden. He maintains the Angiosperm Phylogeny Website: http:\/\/www.mobot.org\/mobot\/research\/apweb\/ "},{"has_event_date":0,"type":"arnoldia","title":"The Sand Pear-Pyrus pyrifolia","article_sequence":5,"start_page":28,"end_page":29,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25478","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15e896f.jpg","volume":67,"issue_number":4,"year":2010,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"The Sand Pear--Pyrus pyrifolia Peter Del Tredici I n more than thirty years at the Arnold Arboretum, I have observed many trees in our collections. Some have not lived up to expectations, but others have proven themselves over time. One such tree is the specimen of sand pear (Pyrus pyrifolia, accession 7272-C) growing at the back edge of the open meadow below the summit of Bussey Hill, which I consider to be the most beautiful flowering tree in the Arboretum. This tree comes into flower in late April or early May, depending on the weather, and at its peak bloom it shines like a beacon in the early spring landscape. When first glimpsed from Bussey Hill Road, against a backdrop of tall white pines, it looks like a giant white cloud-- an effect that is intensified because no leaves compete with the floral display. The flowers are pure white with crimson anthers, 3 to 3.5 centimeters (1.2 to 1.4 inches) in diameter, and are borne in rounded clusters on slender stalks. In bloom, the tree can be easily spotted from the top of Peters Hill, some 800 meters (2,600 feet) away as the crow flies. It stays in flower for up to a full week, holding up well through all kinds of inclement early spring weather. In fall, the tree's glossy, dark green leaves turn beautiful shades of orange and red. Its hard, round fruits are 3 to 4 centimeters (1.2 to 1.6 inches) in diameter, brown, and covered with pale dots. The fruit has an extremely gritty texture (hence its common name--sand pear) and a puckery aftertaste when bitten into. It's hard to imagine how the delectable Chinese and Japanese \"apple-pears\" in the supermarket were derived from this astringent ancestor. The magnificent sand pear on Bussey Hill stands 16.9 meters (55.4 feet) tall with a spread of 25.7 meters (84.3 feet) and a trunk DBH (diameter at breast height) of 79 centimeters (31.1 inches). Remarkably, it seems never to have suffered any major snow, ice, or wind dam- age--an unexpected observation given its age (101 years) and the exposure of the site where it is growing. Such structural integrity provides a striking contrast to the widely planted but notoriously weak `Bradford' Callery pear (Pyrus calleryana `Bradford'), which shows an all too predictable tendency to split apart in severe storms after about age 20. Were it not for its relatively large, messy fruits, our streets might well have been planted with sand pears instead of Callery pears. The Arboretum's beacon tree was grown from seed collected by E. H. Wilson in the fall of 1907, somewhere in the mountains surrounding the city of Ichang in Hupeh (now Hubei) Province. When Wilson collected the seed he did not give the tree a species name, but noted that the Chinese called it \"tang li tzu.\" At the time, sand pears were classified as Pyrus sinensis, a name which was used mainly to describe cultivated plants with large, edible fruits. Back at the Arboretum, Alfred Rehder decided that Wilson's tree was the wild ancestor of these cultivated trees and, in 1915, proposed the name Pyrus serotina for Wilson's specimens. Taxonomy is ever changeable, though, and in 1926 the Japanese botanist Nakai reduced Rehder's name to synonymy with Pyrus pyrifolia--the name the species now bears. Wilson's sand pear seeds arrived at the Arboretum in April 1908 and germinated in the spring of 1909. Sometime prior to 1918, at least three of the seedlings were planted on the grounds. Remarkably, all three are still alive today--a testament to the toughness and tenacity of the species. Specimen 7272-C is the finest of the three, and it will, I hope, remain a shining spring beacon for Arboretum visitors for many years to come. Peter Del Tredici is a Senior Research Scientist at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23417","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160ab6e.jpg","title":"2010-67-4","volume":67,"issue_number":4,"year":2010,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"An Excerpt From Wilson's China: A Century On","article_sequence":1,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25471","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15eb726.jpg","volume":67,"issue_number":3,"year":2010,"series":null,"season":null,"authors":"Flanagan, Mark; Kirkham, Tony","article_content":"An Excerpt From Wilson's China: A Century On Mark Flanagan and Tony Kirkham Editor 's N ote : Ernest H enr y W ilson was one of the most intrepid and productive plant hunters of his era--the beginning of the twentieth century. His collecting trips to China--first for Veitch Nurseries in England and then on behalf of the Arnold Arboretum--resulted in an extraordinary stream of new plants to the West. Arboretum director Charles Sprague Sargent instructed Wilson to thoroughly document his 19071908 and 1910 expeditions with photographs; these striking images still reside in the Arboretum's archives. Mark Flanagan, Keeper of the Gardens at Windsor Great Park , and Tony Kirkham, Head of the Arboretum at Kew, are modern-day plant hunters, having traveled and collected extensively in eastern Asia. Admirers of Wilson, they plotted a journey to retrace his footsteps in Sichuan, China. Using Wilson's expedition photographs as a guide, they were able to capture views of some of the very same locations and even plants that Wilson saw a century ago. Their book pays homage to Wilson and provides a fascinating \"then-and-now\" glimpse of China's landscape. The following is an excerpt from Chapter 3, \"Mystery Towers of Danba.\" Tatien-lu is a small and filthy dirty place, it boasts a large mixed population of Chinese and Tibetans. Being on the highway from Pekin to Lhasa, officials are constantly passing and re-passing. This makes it a highly important place, both politically and commercially. Although Batang, 18 days journey to the west, is the actual frontier town, Tatien-lu is really the gate of Tibet.1 ilson's accurate but rather unflattering description of Kangding was penned at the conclusion of his first visit to the town in 1903. Wilson had made the journey to Kangding on the instructions of the Veitch nurseries who wished to add a very special plant, the lampshade poppy (Meconopsis integrifolia), to their nursery catalogue. \"Messrs. Veitch despatched me on this second, and very costly, journey to the Tibetan border for the sole purpose of discovering and introducing this, the most gorgeous alpine plant extant,\" recorded Wilson.2 Kangding still has a frontier town feel about it and is inalienably a Tibetan place. It also remains a very important staging post on the road that leads westward into Tibet. This road was one of the great highways constructed during imperial times to hold the Celestial Empire together. But Chinese writ did not extend very far. Indeed the country around Kangding remained lawless and untamed until very recent times. Historically the area was known as Kham and its inhabitants, the Khampas, were much feared for their ferocity and war-like demeanor. In truth, Khampa was a collective noun as the area was home to a very diverse group of related, though distinct, peoples. For a thousand years, after the collapse of the vast Tibetan Empire in the ninth century, Kham remained unconquered and unconquerable, its peoples engaged in ceaseless internecine W Wilson's China 3 All PHoToS ARE By THE AuTHoRS unlESS oTHERWISE InDICATED The authors, Mark Flanagan (left) and Tony Kirkham (right), surrounded by prayer flags at the Ya-jia Pass. conflict as petty warrior chieftains battled for supremacy. Banditry was an accepted means of acquiring wealth and position. The various groups of ethnic peoples he encountered fascinated Wilson and he wrote about them extensively. His understanding was gained both first hand and through studying the work of contemporary ethnographers, though this understanding was far from exact. For example, he used the Chinese generic, and derogatory, name \"Sifan\" (western barbarian) to describe the tribe now identified as the Qiang, one of the 56 official ethnic peoples in China. To the enquiring Edwardian mind the alien culture and manners of the various peoples, particularly their peculiar (and supposedly immoral) sexual liaisons in which both polyandry and polygamy were commonplace, was of abiding interest. Their relative lack of sophistication also appealed to Wilson, suggesting to him a oneness with their environment that he found endearing. The eventual subjugation of the Kham region in what the Chinese called the \"peaceful liberation of Tibet\" finally ended the brigandry and general lawlessness in the 1950s. Eastern Kham formally became part of Sichuan Province and Western Kham formed a large part of the Xizang Autonomous Region. Despite this the Khampas retain their individuality by virtue of their strong culture and association with their land. In travelling this country it is impossible not to be impressed by their proud bearing and independent mien and it is easy to 4 Arnoldia 67\/3 understand how they struck fear into the hearts of friend and foe alike. Despite the suppression of banditry, in recent years occasional acts of violence against foreigners still occur when travelers are held-up by groups of armed local men: old habits die hard.3 Through the 1980s and 1990s the Kew expeditions to this part of China employed the services of an armed Chinese policeman, lao liu, as a precaution against unwanted local attention, though he never drew a weapon in anger! Wilson's quest for the lampshade poppy was, therefore, into territory that he knew little of and amongst people with whom, at that stage, he was largely unfamiliar. not only was the territory unfamiliar, it was built on a grand scale. The Da Xue Mountains into which Wilson was travelling, together with the neighboring ranges form part of the vast, complex Hengduan Shan, the eastern extension of the Himalaya. They were created at the same time but, due to the shearing effect involved when the landmass of India collided with the Asian continent, they incline northsouth. These mountains, eroded by monsoon-swollen rivers--the Jinsha, yalong, Dadu Finding the lampshade poppy (Meconopsis integand Min--form an enormous convoluted mass rifolia subspecies integrifolia) was the principal of peaks, ridges and spurs with deep, sheer-sided objective of Wilson's second trip to China. valleys. The range climaxes at the summit of the mighty Gongga Shan, which at 7,556 meters is Sichuan's highest mountain by some way. Joseph Rock brought Gongga Shan to the attention of the West in 1930, when he infamously over-estimated its height, erroneously claiming it to be higher than Mount Everest.4 Wilson would be travelling at far higher elevations and over much more demanding terrain than he had experienced in his first trip to the more modest hills and valleys of Hubei. He was not alone, however. It is intriguing that Wilson rarely mentions any western companions in his writings, let alone provides any details of their backgrounds and occupations. This time, as he prepared to find the lampshade poppy, he was accompanied by an experienced traveler. on July 16, 1903, he started out for the mountains: \"on this journey I was accompanied by Mr. Edgar of the China Inland Mission, in whom I found a delightful companion . . . leaving by the South Gate we followed the main road to lhasa--a broad, well-paved road.\"5 [...] The main road to lhasa was a well-travelled highway but not one that Wilson would remain on for long. It quickly rises to the Zheduo Pass, which today is still the most commonly taken route into Tibet from western Sichuan. on the flanks just below this pass the lampshade poppy can be easily found and many writers have assumed that this is where Wilson gathered his first plants. But the Wilson's China 5 Zheduo Pass was not Wilson's destination; he was heading for the ya-jia Pass, which followed an alternative and much less-used track to the south. At first the journey was enjoyable and Wilson reveled in his surroundings: \"Our road was through lovely grassy country, with a steady rise. A wealth of many colored herbs enlivened our path,\" and, \"we continued through similar country, with a fine snow-clad peak straight in front of us and another to our left.\" Soon, however, the going became much tougher, and heavy rain fell as they reconnoitered the mountainsides close to their overnight stopping point. The altitude had a detrimental effect on his coolies and all endured a miserable night. our journey up to the ya-jia Pass was rather more comfortable. The road was well surfaced right to the top, though the occasional small landslip had to be carefully negotiated by the vehicles. As Wilson suggested, snow-clad peaks were visible all around and we were fortunate to have fine weather in which to appreciate them. looking back, a stunning range of mountains could be seen to the north-east beyond Kangding--the lian lua Shan (lotus Flower Mountains)--no doubt one of the views enjoyed by Wilson during his own ascent 103 years before. Ahead the scene was much less promising with dark clouds scudding across the sky alternately revealing and concealing the mountain tops and providing tantalizing glimpses of the pass. A view to distant Gongga Shan, showing its distinctive peak. 6 Arnoldia 67\/3 Wilson's miserable end to the day was compounded during the night: Having at length got rid of our soaked garments--a difficult enough task under the circumstances--we eventually got between the blankets. No sooner had I lay down than a drip came a spot of rain into my eye: I turned over and drip came another into my ear. I twisted this way and that way, but there was no escape. Like evil genii these rain-drops pursued me turn which way I would. I could not move my bed, since this was longer than the tent was broad, and my feet already exposed, and we sorely afraid the whole thing might collapse, it being anything but secure . . . About 4 a.m. our firewood gave out and things assumed a very dismal aspect. However, all things have an end; day at length dawned and all were devoutly thankful . . . With what fire remained we managed to boil some water and make some tea. We breakfasted on ship's biscuits and cheese and felt none the worse for the night's experience.8 Wilson was, above all things, a fatalist. The rain stopped and Wilson and Edgar prepared for the day's work, during which they hoped to find the lampshade poppy. A farmhouse, one thousand feet below their overnight position, was commandeered and their cook, who was suffering from severe altitude sickness, was taken down to recuperate. The journey to the pass began at 7.a.m. and after some initial rain showers continued on in sunshine. The alpine flowers captivated Wilson; in early summer these Chinese mountains are amongst nature's most exquisite natural gardens. Tony and I arrived at the peak of the display and we left the vehicles three or four hundred meters below the pass and proceeded on foot. By the roadside, a braided mountain stream provided ample moisture and it was in this sodden turf that the greatest diversity could be found. \"I wish I had the ability to describe this floral paradise with all its glories, but this is beyond me,\" wrote Wilson.9 I certainly won't try where Wilson failed and hope that the images reproduced The flora of the moist ground below the Ya-jia Pass is replete with a wonderful array of colorful flowers, including Rheum alexandre (left) and Primula secundiflora (right) Wilson's China 7 on these pages will give the reader a hint of the individual and collective beauty of these mountain flowers, many of which have become firm garden favorites amongst discerning growers. We followed Wilson's and Edgar's route to the pass knowing that at any time the lampshade poppy would appear. our experience was almost exactly as theirs had been: At 11,000 feet I came across the first plant of Meconopsis integrifolia! It was growing amongst scrub and was past flowering. I am not going to attempt to record the feelings which possessed me on first beholding the object of my quest to these wild regions . . . I had travelled some 13,000 miles in five and a half months and to be successful in attaining this first part of my mission in such a short time was a significant reward for all the difficulties and hardships experienced en route.10 The lampshade poppy is a monocarpic species, dying after flowering, but it produces ample seed and has proved to be relatively amenable in cultivation particularly in the cool summer climate of Scotland.11 Wilson's plant became an instant success, flowering in its first season in the Veitch nursery and persisting for many years.12 All the recent trips to this and neighboring parts of China have reinforced its presence in cultivation and it is not unusual to see this plant flowering in northern gardens. In cultivation it has also produced several attractive hybrids with other Asiatic species such as M. beamishii (M. integrifolia M. grandis) and M. finlayorum (M. integrifolia M. quintuplinervia). In recent years botanical opinion, particularly that of Dr. Chris Grey-Wilson, has suggested that this variable plant is easily divisible into two distinct entities-- M. integrifolia and M. pseudointegrifolia, the latter a plant with nodding and more open flowers, quite distinct from Wilson's plants that have globular and more upright flowers.13 After this first plant the mountainsides began to reveal a veritable cornucopia of poppies. Wilson recorded that \"as we continued the ascent, Meconopsis integrifolia became more and more abundant. At 12,000 feet and upwards, miles and miles of the alpine meadows were covered with this plant, but only a few late flowers remained.\"14 Being a month earlier Tony and I caught every plant in full flower, the sun-disk blooms swaying in the mountain breeze, flaunting their wares for any passing bees. our climb continued in deteriorating conditions until we reached the pass at nearly 4,000 meters. Wilson tells us of the fear that the ya-jia Pass engendered amongst his Chinese followers who were not, by nature or inclination, mountain people: \"this Ya-kia pass enjoys an unenviable reputation, and is much dreaded on account of its asphyxiating winds. It is said to be the only pass in the neighborhood which `stops peoples' breath'.\"15 on reaching the pass we were forced to concur, for though it was June 17, the temperature hovered around freezing point and a biting wind blew from the bleak Tibetan Plateau to the west, bringing pulses of sleet in its wake. Despite this we were thrilled to take an image at almost exactly the same location as Wilson had when he re-visited the pass on 19 July 1908. Wilson stayed a second night on the mountain, this time in the more salubrious surroundings of the farmhouse that had been commandeered for the use of 8 Arnoldia 67\/3 ARCHIVES oF THE ARnolD ARBoRETuM The caption for this 1908 Wilson photograph reads: Tachien-lu, near, W. Szechu'an. The Ya-chia-k'an snows and alpine regions clothed with dwarf junipers and rhododendrons. 13,000 feet. July 19, 1908. his party. This proved to be a clean, dry, cozy dwelling, and to add further to their good fortune his cook was quite recovered and prepared a hot meal for the team. on the next day of our trip a most intriguing incident occurred, something that caught me quite by surprise. I have already mentioned Gongga Shan, the giant peak that dominates the Da Xue Shan range. It is the highest mountain in Asia outside of the main Himalayan chain and it exerts a baleful influence. numerous glaciers grind their way down its flanks and such is its size that it generates its own climatic conditions over a sizeable swathe of the surrounding country. This mountain has always fascinated me. Despite dominating the area it is frequently covered in cloud: I have journeyed to five key vantage points-- east, northeast, west, and south of the peak--and been disappointed to find a shroud-covered summit every time. In all his writing Wilson never mentions this mountain either as Gongga Shan or Minya Konka, its Tibetan name. How can this be? During his visits to China he spent many months in the Da Xue Wilson's China 9 A current view of the barren and desolate Ya-jia Pass, unchanged since Wilson and Edgar first came here. Shan, surely he must have heard some local reference to the peak or glimpsed some distant view? Given his silence on the matter the obvious conclusion was that he was also unlucky and never had a clear view of the summit nor did he hear mention of it amongst the local people. As Tony and I wandered the lonely slopes around the ya-jia Pass I pondered this matter, knowing that the giant mountain lay to the southwest of our position. All around us were shattered and snow-clad peaks. It would have taken a strenuous hike into the higher reaches to breast these in order to provide an unencumbered view to the southwest, and time didn't allow this opportunity. In the warmth and comfort of our 4x4 as we took the road back to Kangding, I re-read Wilson's account of his first journey to the ya-jia Pass, particularly the second day of his visit. one paragraph leapt from the page. Although I had pored over all Wilson's writings for the best part of the previous 18 months, the significance of the words had, until now, escaped me: The moraine in front of us terminated in tremendous fields of ice, glaciers of a virgin peak, 21,000 feet high. The sun shone brilliantly and we got a magnificent view of the surrounding mountains. South, south-west of us lay a gigantic peak, several thousand feet higher than the one mentioned; its summit crowned with snowfields of enormous size.16 10 Arnoldia 67\/3 Gongga Shan? Surely. The following day it was time to move on to the next phase of our journey. Wilson's first trip in the employ of the Arnold Arboretum, his third visit to China, took place between 1907 and 1909. Released from the economic shackles imposed by the Veitch nursery he could take a much more expansive view of his activities. The patrician director of the Arnold Arboretum, Charles Sprague Sargent, encouraged Wilson not only to \"science-up\" his work--more emphasis on herbarium specimens and greater attention to field notes--he also insisted that a comprehensive photographic record of the journeys be produced. In a letter to Wilson dated 6 november 1906, a copy of which can be found in the Wilson archive at the Arnold Arboretum, Sargent explains: I write to remind you of the very great importance of the photographic business in your new journey. A good set of photographs are really about as important as anything you can bring back with you. I hope therefore you will not fail to provide yourself with the very best possible instrument irrespective of cost. The hardy and ubiquitous Rhododendron prezwalskii covers huge areas of the high mountains above Kangding. Wilson's China 11 Sargent's prescience not only provided us with an excellent series of images of plants and landscapes, which were later published by the Arnold Arboretum, but also a snapshot of Imperial China right at the end of its long history; within a year of Wilson's departure China was effectively a republic. Thus equipped and instructed Wilson arrived at yichang, his old base on the yangtze River, in February 1907 for what was The local flora is occasionally put to a novel use; this Tibetan girl is using an to be his most successful Incarvillea flower as a kazoo. trip, a trip that cemented his reputation as the foremost collector of his generation. I have long felt that the second year of this expedition, 1908, was also his most interesting and productive and in following in Wilson's footsteps I was especially keen to emulate some of his travels during that year. From Kangding we had the opportunity to retrace Wilson's journey of JuneAugust 1908 when he travelled between Dujiangyan (Kuan Hsien) and Kangding, though we would travel it in reverse. Interestingly, Wilson himself was following an earlier traveler--Sir Alexander Hosie--as he tells us: During the summer of 1908, when in Chengtu, I determined upon a journey to Tachienlu. Previously, in 1903 and again in 1904, I had visited this town by three different routes. This time I decided upon following the road leading from Kuan Hsien via Monkong Ting and Romi Chango. The only published account of this route that I had knowledge of is a report by Mr. (now Sir) Alexander Hosie, erstwhile HBM's Consul-General at Chengtu, who returned over this road in October 1904.17 The account Wilson refers to, \"Journey to the Eastern Frontier of Thibet\", was published as a Parliamentary Report and presented to Parliament in 1905. Hosie took the same direction as Tony and I would, east from Kangding, which he left on 10 october 1904. This was by no means a regular or accepted highway and that is what interested Wilson: \"what I saw of the forests and mountain scenery, together with the quantity and variety of the plants discovered and collected, abundantly repaid me for the hardships experienced.\"18 My hope was that we could also experience some of this scenery and plant diversity. But could we retrace the route and match some of the many images that Wilson had taken on this journey? Things began disappointingly. Wilson had travelled on foot on the east side of the Da Xue Shan between Kangding and the village of Hsin-tientsze and even 12 Arnoldia 67\/3 The new temple at Tagong with the massive bulwark of the Da Pao Shan behind. today there is no suitable road for motorized vehicles. This meant that we would have to drive up the west side of the range before rejoining Wilson's route beyond Hsin-tientsze. Fortunately, apart from stunning views of some of the snowclad peaks and a range of hot springs at Je-shuit'ang, it seemed we would miss nothing of great import. no images of particular interest record this section of the journey. We left Kangding taking the road up and over the Zheduo Pass. In the sunlight the roadsides were bright with wild flowers, many of a striking nature, including the large flowered but short-statured Tibetan lady's slipper orchid (Cypripedium tibeticum) with large maroon pouches. At the pass we had a something of a shock. Having been at this lonely spot in 2001 we were dismayed to find that things were much changed. A wooden belvedere had been built about 150 meters above the pass, reached by a flight of steps, and another building was under construction nearby. no doubt these developments are underpinned by good intentions, this spot is very much on the tourist route, but it somehow seems quite inappropriate to despoil these pristine alpine areas with such frippery. We didn't dwell. on the other side of the pass we got stuck behind an endless convoy of lumbering army trucks, which slowed the pace considerably. one of the positive results of this inconvenience was that we were able to continue to admire the carpet of flowers in the grassy alpine pastures. unlike the valleys to the east, the west Wilson's China 13 side of the Da Xue Shan is quite dry and few trees are to be found. As a result extensive grasslands are a feature and at this time of year they boasted a display worthy of the most colorful flower garden. The turf was studded with gorgeous plants--Incarvillea delavayi, Meconopsis horridula, Lilium lophophorum--odd specimens of Rhododendron capitatum formed hummocky mounds amongst the grass sward and the horizon was an endless, undulating green line. Eventually we turned north leaving the army to continue their procession into Tibet. The road became more and more potholed and uneven as we proceeded. Along the way the solid architecture of Tibet began to dominate, with farm buildings of substantial size and construction. Many are only seasonally occupied as the inhabitants leave in the spring to spend the summers in the high mountain pastures grazing their herds of yak. We passed through the important religious centre of Tagong, dominated by its richly decorated and ornamented temple. Rising in front of us was another range of impressive peaks, the Da Pao Shan (Big Cannon Mountains). This linked us back with Wilson who enjoyed fine weather during the last leg of his journey, albeit on the other side of the range to our position: The view from the summit of the pass far surpassed my wildest dreams. It greatly exceeded anything of its kind that I have seen and would require a far abler pen than mine to describe it adequately. Straight before us, but a little to the right of our view point was an enormous mass of dazzling eternal snow, supposed to be, and I can well believe it, over 22,000 feet high. Beneath the snow and attendant glaciers was a sinister-looking mass of boulders and screes.19 unfortunately for us low clouds obscured the actual summit, though Xiao Zhong told us of that on a recent previous visit he had seen nothing of the mountains at all, so perhaps we were not so unlucky. [...] REFEREnCES 1 12 2 3 4 5 Wilson, E. H. (1906). leaves from my Chinese notebook. Gardeners' Chronicle Vol 39. 17 February. p. 101. Wilson, E. H. (1906). op cit. 24 March. p. 179. \"lonely Planet writer vanishes on trek in Tibet bandit country\". The Times 5 June 2007. Goodman, J. (2006). Joseph F. Rock and His Shangri-La. Caravan Press, Hong Kong. p. 97. Wilson, E. H. (1906). op cit. 3 March. p. 138. [...] Wilson, E. H. (1906). op cit. 3 March. p. 139. Wilson, E. H. (1906). op cit. 17 March. p. 166. Wilson, E. H. (1906). op cit. 24 March. p. 179. Grey-Wilson, C. (1993). Poppies: A guide to the Poppy Family in the Wild and in Cultivation. london. B T Batsford. p. 186. 13 14 15 16 17 18 19 8 9 10 11 Gardeners' Chronicle Vol 36. 1 october. 1904. p. 240. Grey-Wilson, C. (1996). The yellow Poppywort and its Allies. The New Plantsman 3 (1) 2239. Wilson, E. H. (1906). op cit. 24 March. p. 179. Wilson, E. H. (1906). op cit. 17 March. p. 165. Wilson, E. H. (1906). op cit. 24 March. p. 180. Wilson, E. H. (1913). A Naturalist in Western China. Methuen and Co. london. Vol. 1. p 170. Wilson, E. H. (1913). op cit. Wilson, E. H. (1913). op cit. Vol 1. p. 200. Wilson's China: A Century On Mark Flanagan and Tony Kirkham. Kew Publishing. 2009. 256 pages. ISBn-13: 978-1842463949 "},{"has_event_date":0,"type":"arnoldia","title":"Aronia: Native Shrubs With Untapped Potential","article_sequence":2,"start_page":14,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25472","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15eb76b.jpg","volume":67,"issue_number":3,"year":2010,"series":null,"season":null,"authors":"Brand, Mark","article_content":"Aronia: Native Shrubs With Untapped Potential Mark Brand T he genus Aronia is a group of largely overlooked shrubs native to the eastern United States. Aronia species have tremendous potential for use as ornamental landscape plants and as an edible fruit crop. One thing that has held back consumer acceptance of Aronia is the unfortunate common name chokeberry--a name unlikely to endear a plant to consumers. The name chokeberry may have been given to Aronia because people have observed that the berries are initially overlooked by birds and are only taken later in the winter when they are the last fruits remaining. The strong tannin flavor of chokeberry fruits may seem to be the reason why birds avoid the fruits, but ornithologists point out that it may actually be the relatively low protein content of the fruits compared to other fruits that are more readily taken by birds. I am always working to enlighten people about Aronia and in doing so I have found that confusion abounds when it comes to chokeberry and chokecherry. I regularly have people tell me they are familiar with chokeberry, only to find out that they meant chokecherry (Prunus virginiana). Aronia is one of the best kept plant secrets around--surprising since this genus is as complex and interesting as it is useful. Aronia Species and Their Characteristics Chokeberries are in the Rosaceae and are multistemmed, deciduous shrubs. They readily form rhizomes and can sucker to form small colonies in a non-aggressive manner. Two species All phOTOS ARe by The AUThOR UnleSS OTheRwISe IndICATed Red chokeberry's striking fruit display lasts several months. Aronia 15 The leaves of red chokeberry (seen here) are pubescent on the lower surface while black chokeberry leaves lack pubescence. species, A. prunifolia (purple chokeberry), is generally recognized as having purple-black fruits and amounts of pubescence intermediate between the red and black species. In my observation, the amount of pubescence on plants that could be considered A. prunifolia can range from moderate to heavy. Table 1 summarizes some of the characteristics that can be used to try to differentiate red from black chokeberry. Speciation within the Aronia genus is far from clear cut and more research needs to be conducted to determine if A. prunifolia is a hybrid between A. arbutifolia and A. melanocarpa or should be considered as part of the A. melanocarpa species. (See the taxonomy sidebar for more information on Aronia speciation). The red chokeberry grows 6 to 10 feet (1.8 to 3 meters) tall and 3 to 5 feet (.9 to 1.5 meters) Though not long lasting, red chokeberry's flowers are attractive in the spring. Red chokeberry has an upright growth habit. of Aronia are generally recognized: A. arbutifolia (red chokeberry) and A. melanocarpa (black chokeberry). hardin (1973) suggests that fruit color--red versus black--should be used to differentiate between species. In addition to fruit color, Krussmann (1986) used degree of pubescence on stems, leaves, and inflorescences to distinguish red from black chokeberry. A third Red chokeberry has outstanding red fall foliage color. 16 Arnoldia 67\/3 wide. It is a multi-stemmed shrub with a distinctly upright growth habit. even though the plant suckers and spreads, it can become somewhat leggy and open at the base. Most of the foliage on a mature red chokeberry will be found in the upper half of the plant. Summer foliage of red chokeberry is shiny or flat green above and grayish tomentose below. new growth on stems is also quite pubescent. leaves are obovate or elliptical with a short acuminate tip and marginal serrations. Red chokeberry fall foliage turns a vibrant red crimson or purple red and can be spectacular in sunny locations. even in partly shaded locations the leaves muster a very nice blend of orange and red. In addition to being attractive in the summer and fall, the red chokeberry also flowers in spring, usually in early May in new england. Small white flowers are produced in clusters that are about 1.5 inches (3.8 centimeters) wide and can be so numerous that they cover the canopy surface. The flowers do not last a particularly long time (about the same amount of time as Amelanchier flowers), but they do add early season interest to the plant. perhaps the best part about the flowers is that they give rise to abundant red fruits in late September and early October. The clusters of small (0.25 inch [.64 centimeter] diameter) fruits are quite showy and typically remain firm, glossy, and attractive through december. As stated before, birds rarely strip the fruits from the plants until after they have lost ornamental appeal. The black chokeberry can generally be distinguished from the red chokeberry (when fruit are absent) by the lack of pubescence on stems and leaf undersides. black chokeberries are also shorter than their red-fruited counterparts, attaining a mature height of 4 to 8 feet (1.2 to 2.4 meters). like the red chokeberry, it suckers profusely, but forms dense plants and colonies, rarely appearing very leggy. black chokeberry has outstanding, lustrous, dark green summer foliage that turns a pleasing blend of yellow, orange and red in the fall. while Black chokeberry bears glossy black fruit. Aronia 17 the black chokeberry's autumn foliage display may fall a bit shy of that of its red-fruited relative, it is still superior to many shrubs. Flowers are white, borne in May, and are similar in landscape effectiveness to the red chokeberry. The black fruits, from which A. melanocarpa gets its common name, are shiny and larger (0.3 to 0.5 inch [0.8 to 1.3 centimeters] diameter) than the fruits of A. arbutifolia. Fruits can ripen as early as mid-July, but they primarily ripen dur- Table 1: Comparison of red (Aronia arbutifolia) and black (A. melanocarpa) chokeberry characteristics Red fruit color cherry red fruit relatively small ( 0.3 inch) fruit ripens Sept.Oct. fruit persistent into winter leaves, stems, inflorescences pubescent habit upright, leggy at base found primarily on damp\/wet sites inhabits coastal southeastern U.S. BlaCk fruit color black fruit relatively large ( 0.3 inch) fruit ripens late JulyAug. fruit shrivels and drops leaves, stems, inflorescences glabrous habit rounded, full to base found on both damp\/wet sites and dry sites inhabits northeastern and midwestern U.S. The glabrous foliage of black chokeberry is green in summer and can develop good red to orange and yellow fall color. 18 Arnoldia 67\/3 a powerline cut with sand overlaying moist seeps is home to red chokeberries in North Carolina. ing the month of August. black chokeberries wither soon after ripening and either drop off or persist for a while as \"raisins\" on the plant. A. melanocarpa populations in the upper Midwest typically have more persistent fruit than populations in the northeast. distribution and Habitat The geographical range for Aronia arbutifolia is centered in the southeastern Coastal plain, but it can be found extending out into suitable habitats westward into the Appalachian Mountains. It ranges from eastern Texas to northern Florida and continues up the eastern seaboard. It is common in much of the Carolinas, Virginia, Maryland, and new Jersey. Although it can be found in new england, red chokeberry occurs much less frequently there and is generally found close to the coast. The center of distribution for Aronia melanocarpa is in the northeastern states and the Great lakes region, with range extension into the higher elevations of the Appalachian Mountains. In the Appalachian Mountains and the northeast there is considerable overlap of the red and black chokeberry range. Although the information is somewhat incomplete, A. prunifolia seems to be found throughout much of the black chokeberry range and extends somewhat into the red chokeberry range. Aronia arbutifolia occurs in bogs, swamps, savannahs, lowland woods, the edges of water bodies, moist rocky seeps, and moist pine barrens. A. melanocarpa occurs in similar wet locations, but can also be found growing on sand dunes, dry rocky slopes, dry bluffs and balds, and grassy areas. you will rarely find A. arbutifolia on the same dry rocky bluffs and Aronia 19 One type of black\/purple chokeberry environment in Maine. dunes where A. melanocarpa occurs, but I have found it growing in thin layers of organic duff on the exposed spines of rock balds. A. prunifolia is found in areas similar to A. arbutifolia, but also in somewhat drier clearings. Cultural Conditions Chokeberries are considered to be hardy to USdA hardiness zone 4 and, with proper genotype selection, the red species can exhibit good heat tolerance as well. plants can be grown successfully in partial shade or full sun, but better flowering, fruiting, and fall color occur in full sun situations. both red and black chokeberries seem to tolerate dry or wet soil conditions, even though the red species naturally occurs most often in wet areas. best growth can be expected in moist soils, but soil type is not critical. Transplanting and establishment are easy with chokeberries even when they are given only modest aftercare. like most members of the Rosaceae, Aronia has a seemingly endless list of insects and diseases that could attack it, but the plants rarely seem to be affected by much and are considered relatively carefree. I have found that powdery mildew can hit A. melanocarpa, but it doesn't seem to show up to any degree on A. arbutifolia. lacebug is one insect that I have observed occasionally afflicting black chokeberry growing on hot, dry sites. Aronia Genetics: Ploidy and apomixis published literature states that A. arbutifolia has a 2n number (number of chromosomes in somatic cells) of either 34 or 68 and A. melanocarpa has a 2n number of 34 (darlington and Janaki 1945). At the University of Connecticut I have an Aronia germplasm collection of over 20 Arnoldia 67\/3 This rocky outcropping in the appalachian Mountains in Tennessee is a typical habitat for Aronia. Aronia 21 O Taxonomic Teasers in Aronia ver the years, Aronia has been placed in numerous genera, including Mespilus, Pyrus, Adenorachis, Sorbus, and Photinia by different taxonomic authorities (Robertson et al. 1991). Rehder (1949) and hardin (1973) chose to use the genus Aronia for the chokeberries. In 1991, Robertson et al., placed the chokeberries in the genus Photinia, citing no differences in floral and fruit morphology between plants formally in the genus Aronia and those in Photinia. According to Robertson et al., red chokeberry becomes Photinia pyrifolia, black chokeberry becomes Photinia melanocarpa, and purple chokeberry becomes Photinia floribunda. The USdA plants database (plants.usda.gov) has adopted Photinia as the genus for the chokeberries, but USdA GRIn (www. ars-grin.gov\/) is still allowing Aronia. likewise, in the new 6th edition of Michael dirr's Manual of Woody Landscape Plants, Aronia is still being used for the chokeberries. Until more conclusive genetic studies are undertaken, there will likely be continued uncertainty about the correct genus for the chokeberries. Another point of nomenclatural uncertainty is with Aronia prunifolia. Should it be considered a separate species or be folded into A. melanocarpa or A. arbutifolia as a variety? If it is a separate species, does it have its origins as an interspecific hybrid of A. arbutifolia and A. melanocarpa and should it be designated as Aronia x prunifolia? Most of the evidence seems to suggest that the purple chokeberry is the result of interspecific hybridization between red and black chokeberry. we know from our own hybridization work that it is relatively easy to cross red and black chokeberries and get offspring that are not the result of apomixis. we have pollinated diploid black to tetraploid red and have many purple plants which are triploid. hardin (1973) points out that garden hybrids between red and black have arisen at times and have been referred to as Aronia floribunda. Some have argued that the naturally occurring A. prunifolia is something different from A. floribunda because it can occur outside areas where the red and black chokeberries are sympatric, but this argument is flawed. It does not take into consideration the likely scenario that interspecific red-black hybrids produce viable seeds apomictically. The purple species could arise at the margins of overlap of the red and black species and then spread by seed to regions far beyond each parent species' range. purple chokeberry could also spread vegetatively by rhizomes. Furthermore, purple chokeberries seem to occur in the greatest abundance and have the most within-population variability in areas where both the red and black chokeberries overlap. paper chromatography done in the 1960s on red, black, and purple chokeberry found that purple chokeberry contained the greatest number of unique compounds in comparison to red and black, adding more weight to the theory of hybrid origin (Alston et al. 1965). These arguments, along with the fact that A. prunifolia generally has morphological characteristics (degree of leaf\/stem pubescence, fruit color, fruit ripening date, plant habit) that are intermediate between A. arbutifolia and A. melanocarpa, seem to tip the balance in favor of hybrid origin. One bit of work conducted in the 1970s, at the now closed long Ashton Research Station at the University of bristol, found that the flavone C-glucoside vitexin is restricted to arbutifolia and prunifolia x arbutifolia material and absent from melanocarpa and prunifolia x melanocarpa material. These findings do not support the involvement of A. arbutifolia in the parentage of A. prunifolia (Anon. 1974). 22 Arnoldia 67\/3 100 accessions of black, purple, and red species. So far, based on flow cytometry results, we have not found any diploid (2n=34) red chokeberries. with additional collecting, we hope to find the elusive diploid A. arbutifolia. black chokeberries collected from outside of new england have all been tetraploids (2n=68), while new england black chokeberries have been diploid (2n=34). There are numerous accessions that we believe to be A. prunifolia and these plants are either tetraploid or triploid. There is mounting evidence that suggests that Aronia is capable of producing apomictic seeds (persson hovmalm et al. 2004). These are seeds that develop without fertilization of the egg and are, therefore, clones of the mother plant. This is particularly true of tetraploid and triploid forms of Aronia. Apomictic seed set has been suspected from observations of the homogeneity in cultivated Russian plant material (poplavskaya 1995). In our own nAnCy ROSe breeding work with Aronia at the University of Connecticut, we have found that seedling populations from tetraploid plants are visually identical to the female parent regardless of the ploidy of the pollen used. when we use a diploid female parent, we get segregation within the population. we have also found that triploid Aronia produce fertile seed, even though triploidy typically results in sterility. It is likely that polyploid forms of Aronia have evolved to produce seed through apomixis as a functional manner in which to reproduce. Why the Interest in Aronia? The future is particularly bright for Aronia and it will undoubtedly emerge from its relative obscurity to serve as both an important ornamental landscape shrub and as a nutraceutical fruit crop. There is growing interest among gardeners, landscapers, landscape architects, and the general public in making better use of our Chokeberries are versatile ornamental landscape shrubs. a planting of black chokeberry is seen here in fall color. Aronia 23 own native plants, especially when they can serve as alternatives to problematic aggressive and invasive exotic species. In the northeast, the popular winged euonymus or burning bush (Euonymus alatus) has become invasive and has even been banned in Massachusetts and new hampshire. Its main landscape attributes are stunning red fall color, dense habit, and easy culture. Since native highbush blueberry (Vaccinium corymbosum) also has excellent red fall color it is often recommended as a replacement for burning bush, but it is not adapted to many of the landscape sites where burning bush has typically been employed. Aronia can serve as a much better alternative to E. alatus because it is site adaptable in addition to having multi-season interest, including red fall color. To become popular ornamental shrubs, chokeberries simply need a little marketing and perhaps a more appealing common name. In addition to uses as an ornamental, black chokeberry is rapidly gaining momentum as a new small fruit crop for many parts of the United States. The blueberry-sized black fruits produced by Aronia melanocarpa have the highest known levels of antioxidants (anthocyanins and flavonoids) of any temperate fruit, five times higher than cranberry and blueberry, and also contain strong anticancer compounds. Aronia has been widely grown in eastern europe and Russia where the fruits are processed and used in beverages, wine, jelly, and baked goods (Kask 1987). In the United States, Aronia is largely unknown as a fruit crop, but there are no obvious limitations to prevent it from becoming popular here as well, especially given the public's growing interest in functional foods. preliminary work in Iowa, Oregon, wisconsin, and nebraska has demonstrated the viability of Aronia as a fruit crop in many regions, including new england. Aronia berries, while edible as a fresh fruit, are much tastier when the fruits have been processed. They are high in sugar (12 to 20% soluble solids), anthocyanins (560 to 1050 mg\/100 g fresh weight), have a ph of 3.3 to 3.7, and 0.7 to 1.2% titratable acidity (Jeppsson and Johansson 2000; Oszmianski and Sapis 1988). Chokeberries are very suitable for industrial aronia juice products (left to right): aronia blended with acai juice to make a fruit juice drink similar to cranberry cocktail, powdered juice to mix into food or drink as a nutraceutical, and a nutraceutical beverage containing aronia juice. processing since they are not prone to mechanical damage during transport and have low pectin content (Jeppsson 1999). Moreover, chokeberries can be harvested by machine (Gatke and wilke 1991) and there is a long harvest window. Since \"chokeberry juice\" is unlikely to sell, it is usually labeled as \"aronia juice.\" wine red to dark purple in color, it is often blended with other more flavorful juices such as apple, cranberry, grape, and black currant to make popular beverages. Other common uses include jellies and jams, syrup, soft spreads, teas, wine, and flavorings for ice cream and yogurt. Aronia juice is also an excellent colorant. The University of wisconsin-Madison Center for Integrated Systems (Secher 2008) evaluated 13 potential uncommon fruits with sustainability potential. Aronia was chosen as the crop with the greatest potential, beating out currants, gooseberries, and elderberries. low input requirements, high adaptability, high pest resistance, high nutraceutical content, short time to first yield, ease of culture, and high machine harvest potential were given as reasons why Aronia is tops for commercial production potential. 24 Arnoldia 67\/3 The fruit of `Viking'(left) and typical wild-type Aronia melanocarpa (right). `Viking' and `Nero' are the primary cultivars available for fruit production in the United States (Mckay 2001). Both are tetraploid forms (Brand, unpublished data) with large, relatively sweet berries and are the highest producing cultivars currently available (Strik et al. 2003). Breeding and Selection breeding efforts to improve ornamental chokeberries at the University of Connecticut are focused primarily on red chokeberry. There is a need to reduce the plant's stature by half and eliminate its tendency toward legginess. Another goal would be increased fruit size to provide a more impressive display in the fall and early winter. Fall foliage impact can be enhanced by improving leaf retention as the leaves turn red; currently available forms of red chokeberry tend to drop leaves too quickly after coloring. Selections can also be made for resistance to powdery mildew in black chokeberry. So far, improving red chokeberry has been challenging because all of the accessions we have are tetraploids that probably produce apomictic seed. Finding a wild diploid A. arbutifolia could prove to be very useful in breeding this species. polyploidy can also make it more difficult to use mutation breeding on red chokeberry due to the extra sets of chromosomes that can mask incomplete mutations. nonetheless, we have made some progress in developing more compact forms of red chokeberry using chemical mutagens and irradiation. european and Russian breeding efforts to enhance black chokeberry for fruit production have been largely constrained by the highly homogenous gene pool in domesticated Russian plant material. To make progress in this area it is, therefore, necessary to broaden the genetic basis through the introduction of germplasm from native stands (persson hovmalm et al. 2004). There is evidence suggesting that flavonoid content of chokeberries can be increased by incorporating native germplasm that contains higher levels of flavonoids into a breeding program (Sueiro et al. 2006). due to apomixis, diploid forms of black chokeberry are at the core of plant improvement efforts at the University of Connecticut. It is unclear whether tetraploid forms are autotetraploids or allotetraploids. It is possible that commercial Aronia 25 Aronia cultivars like `Viking' could be allotetraploids, since Aronia is known to hybridize with plants in the Rosaceae genus Sorbus. Our primary goal at the University of Connecticut is to improve black chokeberry as a fruit crop by increasing levels of antioxidants and anticancer compounds in the fruits while maintaining or increasing fruit size above that found in current commercial cultivars such as `Viking'. Of course, improving fruit flavor is also important. we are currently trying to unravel the genetics that have given us cultivars like `Viking' and `nero'. Some authorities designate the large fruited black chokeberries as Aronia mitschurini (Strik et al. 2003). In the late 1800s and early 1900s, Russian and eastern european breeders had significant Aronia breeding programs. Ivan Michurin, a Russian plant breeder, produced a plant called `likernaya' that is an intergeneric hybrid between Sorbus aucuparia and Aronia melanocarpa (Kask 1987). It is possible that this intergeneric hybrid, or others like it, were eventually back crossed to A. melanocarpa to give rise to cultivars such as `Viking'. by understanding how `Viking' was created, we hope to re-create superior large-fruited forms for the fledgling domestic chokeberry fruit industry. literature Cited Alston, R. e., h. Rosler, K. naefeh and T. J. Mabry. 1965. hybrid compounds in natural and interspecific hybrids. Proceedings of the National Academy of Science 54:14581465. Anon. 1974. Aronia. long Ashton Research Station, University of bristol: Report 1974. bermudez-Soto, M.J., larrosa, M., Garcia-Cantalejo, J.M., espin, J.C., Tomas-barberan, F.A., & GarciaConesa, M.T. 2007. Up-regulation of tumor suppressor carcinoembryonic antigen-related cell adhesion molecule 1 in human colon cancer Caco-2 cells following repetitive exposure to dietary levels of a polyphenol-rich chokeberry juice. Journal of Nutritional Biochemistry 18: 259271. darlington, C. d. and e. K. Janaki. 1945. Chromosome atlas of cultivated plants. George Allen and Unwin ltd., london. Gatke, R. and K. wilke. 1991. Sind Aronia-busche machinell beerntbar? Gartenbau 38:3738. hardin, J. w. 1973. The enigmatic chokeberries (Aronia, Rosaceae). Torreya 100:178184. Jeppsson, n. 1999. evaluation of black chokeberry, Aronia melanocarpa, germplasm for production of natural food colourants. Acta Horticulturae 484:193198. Jeppsson, n. and R. Johansson. 2000. Changes in fruit quality in black chokeberry (Aronia melanocarpa) during maturation. Journal of Horticultural Science and Biotechnology 75:340345. Kask, K. 1987. large-fruited black chokeberry (Aronia melanocarpa). Fruit Varieties Journal 41:47. Krussmann G. 1986. Cultivated Broad-leaved Trees and Shrubs, 3 volumes, b. T. batsford ltd., london. McKay, S.A. 2001. demand increasing for aronia and elderberry in north America. NY Fruit Quarterly 9:23. Oszmianski, J. and J. C. Sapis. 1988 Anthocyanins in fruits of Aronia melanocarpa (chokeberry). Journal of Food Science 53:12411242. paplavskaya, T. K. 1995. Aronia, its economic significance and current status. In: A program and methods of breeding fruit, small-fruit and nut bearing crops. Oryol, Russia:457459. persson hovmalm, h.A., n. Jeppsson, I. V. bartish and h. nybom. 2004. RApd analysis of diploid and tetraploid populations of Aronia points to different reproductive strategies within the genus. Hereditas 141:301312. Rehder, A. 1949. Bibliography of Cultivated Trees and Shrubs. Arnold Arboretum, harvard University. pp. 261262. Robertson, K.R., J.b. phipps, J.R. Rohrer, and p.G. Smith. 1991. A synopsis of genera in Maloideae (Rosaceae). Systematic Botany 16: 376394. Secher, d. 2008. Fruit with potential for wisconsin farms. http:\/\/www.cias.wisc.edu\/wp-content\/ uploads\/2008\/07\/carandale.pdf. Strik, b., Finn, C. and wrolstad, R. 2003. performance of chokeberry (Aronia melanocarpa) in Oregon, USA. Acta Horticulturae (IShS) 626:439443. Sueiro, l., G. G. yousef, d. Seigler, e. G. de Mejia, M. h. Grace and M. A. lila. 2006. Chemopreventive potencial of flavonoid extracts from plantationbred and wild Aronia melanocarpa (black chokeberry) fruits. Journal of Food Science 71:480488. Mark brand is a professor in the department of plant Science and landscape Architecture at the University of Connecticut. "},{"has_event_date":0,"type":"arnoldia","title":"Forest Farming","article_sequence":3,"start_page":26,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25474","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15e8128.jpg","volume":67,"issue_number":3,"year":2010,"series":null,"season":null,"authors":"Mudge, Ken","article_content":"Forest Farming Ken Mudge NaNcy Rose Many sections of the Northeast have been reforested over the past century. Extensive forest cover is seen in this view from Wachusett Mountain in central Massachusetts. F armers harvest crops from their fields, and loggers harvest trees from their forests, but what do forest farmers harvest? The answer is an eclectic collection of non-timber forest crops like maple syrup, medicinal herbs, fruits, gourmet mushrooms, and nuts. Forest farming is an approach to forest management that combines some of the management practices of conventional forestry with those of farming or gardening to achieve an environmentally and economically sustainable land-use system. It is one of several related practices that fall under the domain of agroforestry--a multidisciplinary approach to agricultural production that achieves diverse, profitable, sustainable land use by integrating trees with non-timber forest crops. While some other agroforestry practices begin with planting young trees that take years to mature, forest farming involves planting nontimber forest crops beneath the canopy of an established forest. In other words, other agroforestry practices bring the forest to the crops, whereas forest farming brings the crops to the forest. In this regard it is helpful to consider the role of forest farming in overall forest man- Forest Farming 27 agement. a forest farm should be designed to emulate as much as possible a natural forest. This includes characteristics of a healthy forest ecosystem such as species diversity, resilience to disturbance, soil health, and a relatively wide tree age distribution. Forest Farming Through the Ages although this article will focus on modern temperate region forest farming, similar practices have been used in tropical regions by indigenous peoples for hundreds of years. In a classic paper from the agroforestry literature, Fernandez et al. (1984) described an agroforestry practice called home gardens, used by the chagga people who live on the slopes of Mt Kilimanjaro. Home gardens are highly integrated, multistory collections of overstory forest trees valued for timber, an intermediate layer of small trees including coffee and banana, and a diverse array of understory herbs and vines used for food and medicine. In North america, during and prior to the seventeenth century, native peoples are known to have planted and managed various food bearing trees including walnuts and peaches, but there is no evidence of deliberate cultivation of useful crops beneath the canopy of established forest. although some types of forest farming and other agroforestry practices have been going on for centuries, the terms \"agroforestry\" and \"forest farming\" are of relatively recent origin. agroforestry--as a concept that recognized the integration of trees, crops, and people--was introduced in 1973 by John Bene, and led to the establishment of the International council for Research in agroforestry (IcRaF) in Nairobi, Kenya which is now the World agroforestry center . It was not until 2000 that the term forest farming was introduced by Hill and Buck (2000) to describe the cultivation of non-timber forest crops beneath an existing tree canopy. one factor contributing to the growing popularity of forest farming in northeastern North america is the gradual increase in the extent of privately owned forest. Using the state of New york as an example, forest cover was at a minimum of about 15% in 1880 because of extensive conversion of forest to farmland. NaNcy Rose Farming or Wildcrafting? although there is no anthropological or archeological evidence that Native americans practiced forest farming per se, unquestionably they were highly skilled at gathering and utilizing wild forest products, including food, medicines, and ceremonial plants. Prospective forest farmers today frequently ask if the collection of wild forest products like edible mushrooms, wild leeks (ramps), ginseng and other medicinals, and decoratives like pine cones for wreaths and vines for basketry can be considered forest farming. These practices, known collectively as wildcrafting, are certainly compatible with forest farming, particularly when done on a sustainable basis, but don't qualify as forest farming (i.e. cultivation) unless they are practiced in combination with deliberate cultivation of nontimber forest crops. Wild-collected morel mushrooms. 28 Arnoldia 67\/3 Reforestation since then has gradually increased forest cover to 65%, and about 60% of that is privately owned. at the same time average parcel size has decreased, making timber extraction a less economically feasible option. This transition from \"industrial\" forestry to \"investment\" forestry by owners who consider forestry a part-time activity or even a hobby makes forest farming an attractive management alternative. Those interested in forest farming include conventional farmers, many of whom have woodlots on their farm, but also private forest owners with non-farm day jobs who want to use their forest productively while preserving or restoring the natural ecosystem. In either case, supplemental income associated with sale of non-timber forest crops can be the deciding factor, although many forest owners pursue forest farming as a source of non-timber forest products for the family, or simply as a source of personal satisfaction. The banana-custard-flavored fruits of pawpaw (Asimina triloba) are a potential forest farming product. Becoming a Forest Farmer Before starting, the forest owner should consider site issues beyond just \"can I grow ginseng (and\/or any other crop) at this site?\" a successful forest farm should be seen as an integrated agro-ecosystem that satisfies the owners goals while sustaining ecosystem components including soil, water, trees, and wildlife. a forest farm often begins with a more or less natural (unmanaged) stand of trees. To make it suitable for forest farming, some degree of management is necessary, including management of the forest light environment. For the medicinal herb ginseng (Panax quinquefolius), up to 70% shade is necessary, and for mushrooms like shiitake (Lentinula edodes), the more shade the better. on the other hand, fruit crops like blackberries (Rubus spp.) and pawpaw (Asimina triloba) perform best under moderate shade. Light management practices include pruning, selective tree removal, utilizing natural gaps in the forest canopy, and planting less shade-tolerant crops along the forest perimeter. observing the natural distribution of wild relatives can inform decisions about appropriate placement of candidate non-timber forest crops. For example, wild brambles like black raspberry and blackberry typically occur along the ecotone (interface) between field and forest. attempts to grow cultivated brambles beneath a dense forest canopy rather than along its perimeter generally have not been successful. While light is an environmental factor that can be managed, others, including soil pH, fertility, slope, and water availability, cannot be modified in forests as easily as in field agriculture. Irrigation, fertilization, and modification of soil pH are not realistic management options in most forest situations. It makes more sense to select crops that are naturally well-suited to the site characteristics. Types of Non-Timber Forest Crops There are three major categories of non-timber forest crops used in forest farming: medicinal, food, and ornamental. Ginseng and mushrooms, in the medicinal and food categories respectively, have the greatest proven income potential but there are others in each category well worth considering (see chamberlin et al. 2009). While cultivation of ornamentals in forest farming systems is less frequently practiced it does have considerable potential. In traditional farming, crop diversification was considered insurance against the failure of any one crop. species and temporal diversity are characteristics of a natural forest ecosystem which forest farms ideally should seek to emulate. For example, in a forest farm growing mushrooms, maple products, and ginseng, diversification makes good sense from a production standpoint; maple is tapped yearly, mushrooms yield a harvest for several years, while ginseng will take eight years to mature. In this NaNcy Rose Forest Farming 29 KeN MUdGe case, the maples also provide shade for both other crops and additional calcium (in fallen leaves) needed by the ginseng. Medicinals The forests of North america have been repositories for a wide range of herbs and other plants and mushrooms gathered for use in traditional medicine. The pallet of medicinal plants collected and used by Native americans and others was and is extensive, but only a few of these species are cultivated as non-timber forest crops today. To a very great extent, this is due to economic factors, including lack of markets and concomitantly low potential for income generation. Ginseng and goldenseal (Hydrastis canadensis) are the two medicinals most often cultivated by forest farmers. american ginseng (Panax quinquefolius) is a shade-loving perennial herb that produces a valuable below-ground storage root. It occurs in hardwood forests throughout most of eastern North america. although wild populations have declined somewhat because of harvesting, it is still relatively common. american ginseng has been highly valued in traditional chinese and Korean medicine since it was first exported from North america in the seventeenth century. Most commercial demand today is from china and Korea. Ginseng is valuable as a forest farming product, but potential growers should use Beyfuss's site assessment techniques (see listing under additional Reading) before jumping in. american ginseng is reputed to function in the human body as an \"adaptogen,\" increasing the body's resistance to stress. In traditional chinese medicine, ginseng is said to promote yin energy and have a calming effect. This and other beneficial effects of ginseng have been claimed for centuries, but there is little modern scientific research to substantiate these claims. Nevertheless, ginseng is the most valuable of North american medicinal herbs. The ginseng plant has an unusual growth habit and life cycle that contribute to the relative difficulty and long timeframe involved in cultivating it as a crop. a more typical plant has roots, stem(s) and multiple leaves, each associated with a bud that grows into new branches. Not ginseng. The mature plant consists of a single stalk that American ginseng (Panax quinquefolius). KeN MUdGe Goldenseal (Hydrastis canadensis). looks like a stem, but is in fact a sympodium consisting of the fused petioles (leaf stalks) of its 3 or 4 palmately compound leaves. Ginseng develops a narrow underground rhizome--about the thickness of a good sized-earthworm-- with a bud at one end and one or more tuberous storage roots at the other end. a single flush of aboveground growth emerges from the rhizome bud in the spring while the storage root grows slowly during the summer. The plant's slow growth rate is an adaption to its low light environment and results in forest-cultivated ginseng taking about eight years to mature. The storage root, which looks a little like a branched carrot, is what all the excitement is about. In chinese the word for ginseng means \"man root,\" and the more it looks like a person (arms and legs) the more valuable it is to traditional asian buyers. The root is the source of 30 Arnoldia 67\/3 KeN MUdGe The branched storage root of ginseng. the pharmacologically active compounds known as ginsenosides. When it comes time to harvest the crop, it can only be sold to a licensed dealer if it is intended for export, as is the case with most american ginseng. The price structure for ginseng is a curious inversion of most other crops. The more intensively it is cultivated, the less it is worth. Large scale, high density ginseng grown in Wisconsin, ontario, and British colombia requires expensive artificial shade structures and considerable amounts of fertilizers and fungicides. The wholesale value of this ginseng is about $25 per pound dry weight as of 2009. as a forest farming crop, ginseng is cultivated either by the \"woods-cultivated\" or the \"wildsimulated\" method. \"Woods-cultivated\" ginseng is grown in raised beds, often amended with organic matter. Its price is about $150 per pound dry weight. The less intensive wildsimulated method involves minimal management--little more than roughing up the ground with a rake, scattering the seed, and coming back to harvest eight years later. It wholesales for about $300 per pound dry weight. This inverse relationship between price and intensity of cultivation extends even to wild-collected (zero cultivation) ginseng which wholesales for $400 to $600 or more per pound dry weight. Food North american forests abound with edible plants. However, their use in forest farming is limited since few of these forest edibles can be grown in sufficient quantity, in a reason- able period of time, and be sold for a reasonable price. Food crops that are most likely to be found in a forest farm include gourmet mushrooms like shiitakes (Lentinula edodes), berries, other fruits such as pawpaw (Asimina triloba), ramps (Allium tricoccum), and tree nuts such as walnuts (Juglans spp.) and hickories (Carya spp.). Forest-cultivated mushrooms deserve serious consideration for those starting a new forest farming venture. Most candidate crops like medicinals, fruits and nuts, or ornamentals require specific site conditions, but since mushrooms are grown on logs they are less dependent on factors like soil moisture, pH, and drainage. Mushrooms are not photosynthetic and therefore can tolerate nearly complete shade, and a shady site is essential to minimize excessive drying of the substrate logs. aside from socioeconomic factors like access to markets, mushrooms can be cultivated almost anywhere as long as there is sufficient shade and a source of substrate logs. Mushrooms are less valuable per pound than ginseng, they are more perishable, and their cultivation requires more labor than ginseng. on the other hand, $8 to $16 per pound fresh weight for shiitake mushrooms is not a bad price when you consider that you can start harvesting mushrooms in as little as one year after log inoculation, and continue harvesting from the same log for 3 to 5 years. By contrast, once a ginseng root is harvested, the plant is gone. other mushrooms including lion's mane (Hericium spp.), oyster (Pleurotis spp.), and hen of the woods (or maitake) (Griffola frondosa) can be cultivated under forest farming conditions but cultivation strategies are not yet as well worked out as for shiitake. People curious about mushroom cultivation often ask about the ultra-valuable truffles (Tuber spp.), which can sell for nearly $1000 per pound, and valuable morels (Morchella spp.). It is probably best to discourage all but the bravest and most patient entrepreneurs from investing time or money in cultivating truffles--many have tried to grow them but very few have succeeded. similarly, there are very few who have successfully grown morels, which are best left to wildcrafting. By far the most reliable mushroom for forest cultivation, and for which there is the greatest commercial demand, is shiitake. While log- Forest Farming 31 KeN MUdGe Lion's mane (Hericium spp.) and oyster mushrooms (Pleurotis spp.) can be cultivated in forest farms, though not as reliably as shiitake. KeN MUdGe Forest-farmed shiitake mushrooms ready for harvest. grown shiitake can be a reliable non-timber forest crop for forest farming, it is worth pointing out that most shiitake mushrooms available to the public are grown indoors on artificial (sawdust) logs, in expensive climate-controlled rooms. The quality of these artificially cultivated shiitake is generally considered inferior to log-grown shiitake mushrooms. shiitake is a primary saprophytic fungus that derives its nourishment from dead organic matter; in forest farming, freshly cut logs provide the substrate. The goal of shiitake cultivation is to facilitate the decay of the log through a process by which the fungal mycelium (aggregated fungal strands, or hyphae) enzymatically digests the wood by degrading lignin and cellulose, producing carbon dioxide, water (products of respiration), and the energy necessary for the fungus to assimilate the remaining carbon into new mycelium and reproductive structures, that is, the mushrooms. since shiitake competes poorly with other fungi, the substrate logs for production must be freshly cut and lack competing decay organisms. Freshly cut logs also provide the high moisture content necessary for fungal colonization after inoculation. a number of deciduous hardwood tree species make good shiitake substrate. oaks (Quercus spp.) are considered the gold standard in the Northeast but sugar maple (Acer saccharum) also rates highly. Tulip poplar (Liriodendron tulipifera) is particularly desirable further south. conventional wisdom says that species with tight bark that helps maintain a high moisture content make the best shiitake bolts, yet our research has shown that aspen, which maintains a higher moisture content than oak, beech, or red maple, is the poorest with respect to shiitake production. Interestingly, in our tests ironwood, or american hornbeam (Carpinus caroliniana) outperformed red oak. cut logs are then inoculated with spawn, a pure culture of shiitake fungal mycelium (not spores) grown on sawdust or other substrate which is introduced into holes drilled in the logs. Production of uncontaminated spawn must be done under sterile laboratory conditions, so most growers purchase their spawn from commercial producers. an approximately $20 bag of spawn inoculates about 2030 logs. once inoculated, the logs are transferred to the KeN MUdGe 32 Arnoldia 67\/3 P The MacDaniels Nut Grove: A Unique Educational Site racticum in Forest Farming is a multidisciplinary course taught at cornell University. What makes the course unique is not only the subject matter but also the Macdaniels Nut Grove, the outdoor classroom where the course is taught. Both are an outgrowth of the rediscovery, in 2000, of a more than 70-year-old temperate nut tree variety trial established in the 1920s by Professor Lawrence Macdaniels. Not long after dr. Mac (as he was known) retired in 1956, the site was abandoned. soon, it was all but forgotten as it reverted to secondary forest including oak, hickory, maple, cherry, and invasive honeysuckle. Fifty or more years later, the only obvious sign that the seven acre site was once a repository for hickory and walnut clonal varieties was an abundance of graft unions on over 100 of the older trees. When dr. Mac was acquiring and grafting scions (upper portion of a grafted plant) onto understocks (lower portion), little was known about the limits of genetic compatibility between different hickory (Carya) species. Graft unions between genetically compatible scion\/stock combinations like C. x dunbarii (C. laciniosa x C. ovata) grafted onto a shagbark hickory (C. ovata) understock are barely evident today. other combinations like shagbark hickory grafted onto red pignut hickory (C. ovalis) showed extreme bulging and cracking at the graft union, signs of delayed incompatibility. When the site was rediscovered it was easy to recognize it as a nut tree variety trial, but it didn't take long to envision it as an outdoor forest farming classroom for cornell students and members of the community. The course is structured around experiential learning, combining a wide range of outdoor activities with related reading and writing assignments. a field trip to cornell's arnot Teaching and Research Forest lets students learn about ongoing mushroom and ginseng research, A view into the MacDaniels Nut Grove, teaching site for Cornell's including hands-on inoculation of freshly Practicum in Forest Farming. cut logs with shiitake mushroom spawn and digging young ginseng seedlings from a wild-simulated (not wild) ginseng patch. Both the logs and the ginseng are brought back to the Macdaniels Nut Grove for further learning activities. additional activities include soil analysis, vegetation inventory, site indexing to gauge suitability for a given tree species, and forest stand improvement. data collected in each of these categories contribute to an ongoing GPs database of the Macdaniels Nut Grove. as an integrator of all these hands-on activities, student groups work on a final project involving a systematic permaculture-inspired approach to site assessment and design using an explicit set of \"forest owner\" goals developed by each (A) shows an incompatible graft union (Carya ovata on C. ovalis) group in order to synthesize a final design. and (B) shows a compatible graft union (C. x dunbarii on C. ovata). KeN MUdGe KeN MUdGe Forest Farming 33 KeN MUdGe KeN MUdGe thrive in full sun are not well suited for forest farming, but there is plenty of demand for shade-tolerant garden perennials including hostas, daylilies, ferns, heucheras, trilliums, astilbes, and hellebores. Note that some of these are forest wildflowers, but in forest farming the emphasis is on deliberate cultivation and conservation of wild plants so plants are grown, not wild collected. Two basic nursery production systems are adaptable to forest farming--field (in the ground) and container. Field production can have lower costs but harvest and sale of plants is seasonally limited. conA laying yard of logs inoculated with shiitake mushroom spawn. tainer production makes it easier to harvest, move, and sell plants, but has drawbacks such as wind throw, low and high temperature stress to root systems, and higher water needs from restricted rooting volume. a newer method, pot-in-pot production, solves many of these container problems and may be well suited for forest farming. This method uses an empty socket pot buried in the ground so that its rim is about even with the soil surface. a second pot containing the crop plant slides down into the socket pot, so its soil line is at about the natural ground line. This prevents wind throw, allows for easy, multi-season harvest, moderates soil temperature These hostas are being forest-farmed in a pot-in-pot nursery production bed. within the crop pot, and reduces Netting prevents deer browsing. water use. The latter is particularly laying yard where they will incubate while the useful in most forest farming systems where fungal mycelium colonizes the log. The layirrigation is usually not readily available. ing yard must be well shaded (about 80%) year another category of ornamentals for forest round to minimize moisture loss from the logs. farm production includes cut stems, flowers, Production of mushrooms for harvest typically cones, and vines that are harvested from intact occurs about a year after inoculation. plants that remain alive, ready to produce subsequent crops. although collection of cones, Ornamentals boughs, bark, etc., is more closely allied with The North american public spends a considwildcrafting than forest farming, the distinction erable amount of money on plants used as becomes blurred if a wild plant is deliberately ornamentals, either in gardens or for decoramanaged to produce a continuous supply of cut tive purposes around the home (e.g. cut flowproducts. Woody florals--ornamental stems ers). ornamental plants like roses or lilacs that of woody shrubs such as red-twig dogwood or 34 Arnoldia 67\/3 NaNcy Rose Red-twig (or red osier) dogwood (Cornus sericea) can be grown as a woody floral for its colorful winter stems. Forest Farming 35 corkscrew willow used in floral design--are another potential crop, though most woody species in this category would be best suited for planting along the perimeter of a forest farming site. Woody floral shrubs are generally coppiced (cut back to ground level annually) to generate multiple new shoots for harvest each year. The Future of Forest Farming Forest farming is not yet a widespread approach to, or component of, forest management, but it has great potential for wider adoption as forest owners look for alternatives to either management of the forest for timber extraction or no management at all. development of a successful forest farming design should include an integration of the explicit goals of the forest owner with a systematic assessment of both biophysical and anthropogenic characteristics of the site. Ginseng, maple syrup, and forest-cultivated mushrooms have the most reliable track record, and are potentially more profitable than other non-timber forest crops. others like minor fruits and ornamentals have considerable potential for exploration and development. Additional Reading Beyfuss, R. Visual site assessment (for ginseng cultivation) http:\/\/scnyat.cce.cornell.edu\/forestfarming\/ ginsengvisualassmt.pdf chamberlin, J.L., d. Mitchell, T. Brigham, T. Hobby, L. Zabek, J. davis. 2009, Forest Farming Practices, ch. 9, In: North American Agroforestry, An Integrated Science and Practice, 2nd ed., edited by H.e. Garrett. american society of agronomy Press, Madison, WI. Fernandez, e.c.M., a. oktingata, and J. Maghembe. 1984. The chagga homegardens: a multistoried agroforestry cropping system on Mt. Kilimanjaro (Northern Tanzania). Agroforestry Systems, 2: 7386. Hill, d.B. and L.e. Buck. 2000. Forest Farming Practices (ch. 8), In: Garrett, H.e., W.J. Rietveld, and R.F. Fisher (eds.), North American Agroforestry: An Integrated Science and Practice, amer. soc. agronomy, Madison, WI. Jacke, d. and e. Toensmeier. 1996. Edible Forest Gardens, vol. 2, chelsea Green Publisher, White River Junction, VT. Mudge, K.W., 2009, Northeast Forest Mushroom Growers Network http:\/\/mushrooms.cals.cornell.edu\/ Mudge, K.W., L. Buck and P. Treadwell, 2007. How When and Why of Forest Farming http:\/\/www.hort. cornell.edu\/forestfarming\/ Mudge, K.W., 2009. Macdaniels Nut Grove Forest Farm http:\/\/www.hort.cornell.edu\/mng\/ Mudge, K.W. Forest Farming at the Macdaniels Nut Grove, Cornell Plantations Magazine, summer 2004, 59(2):611. World agroforestry center, http:\/\/www.worldagroforestry. org\/af\/index.php Ken Mudge is an associate professor in the department of Horticulture, cornell University, Ithaca, New york. 36673667 U.S. POSTAL SERVICE STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION (Required by 39 U.S.C. 3685) 1. Publication Title: arnoldia. 2. Publication No: 00042633. 3. Filing date: october 14, 2009. 4. Issue Frequency: Quarterly. 5. No. of Issues Published annually: 4. 6. annual subscription Price: $20.00 domestic; $25.00 foreign. 7. complete Mailing address of Known office of Publication: arnold arboretum, 125 arborway, Boston, suffolk county, Ma 021303500. 8. complete Mailing address of Headquarters of General Business office of Publisher: arnold arboretum, 125 arborway, Boston, suffolk county, Ma 021303500. 9. Full Names and complete Mailing address of Publisher, editor, and Managing editor: arnold arboretum, 125 arborway, Boston, suffolk county, Ma 021303500, publisher; Nancy Rose, arnold arboretum, 125 arborway, Boston, Ma 021303500, editor. 10. owner: The arnold arboretum of Harvard University, 125 arborway, Boston, suffolk county, Ma 021303500. 11. Known Bondholders, Mortgagees, and other security Holders owning or Holding 1 Percent or More of Total amount of Bonds, Mortgages, or other securities: none. 12. The purpose, function, and nonprofit status of this organization and the exempt status for federal income tax purposes have not changed during the preceding 12 months. 13. Publication Name: arnoldia. 14. Issue date for circulation data Below: october 8, 2009. 15. extent and Nature of circulation. a. Total No. copies. average No. copies each Issue during Preceding 12 Months: 3,000. actual No. copies of single Issue Published Nearest to Filing date: 3,000. b. Paid and\/or Requested circulation. (1) Paid\/ Requested outside-county Mail subscriptions. average No. copies each Issue during Preceding 12 Months. copies each Issue during Preceding 12 Months: 1,453. No. copies of single Issue Published Nearest to Filing date: 1,421. (2) Paid In-county subscriptions. average No. copies each Issue during Preceding 12 Months. copies each Issue during Preceding 12 Months: 430. No. copies of single Issue Published Nearest to Filing date: 398. (3) sales Through dealers and carriers, street Vendors, and counter sales: none. (4) other classes Mailed Through the UsPs: none. c. Total Paid and\/or Requested circulation. average No. copies each Issue during Preceding 12 Months: 1,883. actual No. copies of single Issue Published Nearest to Filing date: 1,819. d. Free distribution by Mail. average No. copies each Issue during Preceding 12 Months: 108. actual No. copies of single Issue Published Nearest to Filing date: 110. e. Free distribution outside the Mail: average No. copies each Issue during Preceding 12 Months: 500. actual No. copies of single Issue Published Nearest to Filing date: 480. f. Total Free distribution: average No. copies each Issue during Preceding 12 Months: 608. actual No. copies of single Issue Published Nearest to Filing date: 590. g. Total distribution: average No. copies each Issue during Preceding 12 Months: 2,491. actual No. copies of single Issue Published Nearest to Filing date: 2,409. h. copies Not distributed. average No. copies each Issue during Preceding 12 Months: 509. actual No. copies of single Issue Published Nearest to Filing date: 591. i. Total. average No. copies each Issue during Preceding 12 Months: 3,000. actual No. copies of single Issue Published Nearest to Filing date: 3,000. j. Percent Paid and\/or Requested circulation. average No. copies each Issue during Preceding 12 Months: 76%. actual No. copies of single Issue Published Nearest to Filing date: 76%. I certify that all information furnished on this form is true and complete. Nancy Rose, editor. "},{"has_event_date":0,"type":"arnoldia","title":"A Soft Touch: Pinus wallichiana","article_sequence":4,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25473","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15ebb6f.jpg","volume":67,"issue_number":3,"year":2010,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"A Soft Touch: Pinus wallichiana Nancy Rose I f any tree could be described as pettable it would have to be the Himalayan pine (Pinus wallichiana). Its drooping clusters of finetextured, luxuriantly long needles positively invite touching. Think of it as the Afghan hound of the conifer world. Native to the Himalayan Mountains from Afghanistan to Burma, Himalayan pine is fairly common at mid to upper elevations. It grows in valleys and hillsides either alone or with other conifers and deciduous trees including oak, birch, and maple species. Himalayan pine is known to reach 50 meters (about 160 feet) or more in height in its native range. It typically has a straight central trunk and horizontal to slightly drooping branches. In open, cultivated settings it tends to be shorter (perhaps 15 to 25 meters [about 50 to 80 feet]) and develops an attractive domed or haystack shape. When grown like this, with plenty of space, the foliage display is most attractive. The thin, 15 to 20 centimeters (about 6 to 8 inches) long needles are bundled in groups of five and cascade elegantly from the branches like bluish green waterfalls. The pendant cones of Himalayan pine look similar to those of Eastern white pine (Pinus strobus)--light brown at maturity, with flexible scales, and very resinous. Cold hardiness for cultivated Himalayan pine is somewhat variable depending on provenance. The species has grown well in locations in the United States at least as cold as USDA hardiness zone 5 (average annual minimum temperature -20 to -10F [-23 to -29C]). As with some other thin-needled pines, the foliage may suffer winter desiccation damage in windy, exposed sites. The Arnold Arboretum's first accession of Himalayan pine arrived in January 1874 from The Royal Botanic Gardens, Kew (the plant was removed from the Arboretum in 1892 for unknown reasons). The Arboretum's curatorial records show that this accession was received under the name Pinus excelsea, changed to P. nepalensis, back to P. excelsea, then to P. griffithii. These are all synonyms for the nowaccepted name P. wallichiana. The Arboretum holds a number of accessions of Himalayan pine. Currently, the oldest living specimen is a large, handsome 1946 accession (268-46-A) originally received from Karl Sax under the name P. griffithii. It stands 18 meters (59 feet) tall, has a crown spread of 16.4 meters (54 feet), and a trunk diameter of 97.8 centimeters (38.5 inches) (measured below the first limb). This plant is of unknown garden origin, but the Arboretum does have several accessions of documented wild origin as well. The specimen pictured at right (accession 83-94-B) arrived as seed from the Quarryhill Botanic Garden, originally collected in October 1993 during a plant expedition to India. The seeds were collected in the northern Indian state of Himachal Pradesh at an elevation of approximately 2500 meters (8200 feet). Of the four specimens of accession 83-94 planted, this one has performed best, perhaps because of its somewhat protected but adequately sunny location behind the Arboretum's maintenance garage. It currently stands 8.3 meters (27 feet) tall and has a DBH (diameter at breast height) of 17.3 centimeters (6.8 inches). We also have two specimens of another accession, 84-94, which originated from the same 1993 expedition but a different location, this one at an elevation of approximately 2300 meters (7500 feet) in the Great Himalayan National Park. One last wild collected Himalayan pine accession of interest is 1277-61-A, grown from seed collected in November 1961 in the vicinity of Kabul, Afghanistan, at an elevation between 6000 and 7000 feet (about 1800 to 2100 meters). Unfortunately, this specimen is in only fair condition but is a candidate for repropagation because of its unique provenance. Nancy Rose is editor of Arnoldia. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23416","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160ab28.jpg","title":"2010-67-3","volume":67,"issue_number":3,"year":2010,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Crabapples...With No Apologies","article_sequence":2,"start_page":2,"end_page":13,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25468","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15eaf6f.jpg","volume":67,"issue_number":2,"year":2009,"series":null,"season":null,"authors":"Iles, Jeff","article_content":"Crabapples . . . With No Apologies Jeff Iles NANcY ROse An oldie but a goodie, red-flowered `Liset' is still a popular crabapple. O ne of my favorite older horticulture books is a signed copy of Ornamental Crabapples by Arie F. den Boer. Pub lished in 1959 by the American Association of Nurserymen, this little manual was perhaps the first successful attempt at popularizing the various species, varieties, and cultivars of crab apples (those taxa in the genus Malus bearing fruits 2 inches in diameter or smaller). I like the book because it provides a unique glimpse back to an era when selections like `Aldenhamensis', `Almey', and `Dorothea' ruled the nursery sales yards. Those cultivars are rarely seen today but others described in the book, including Malus floribunda, `Liset', `Profusion', and `Red Jade', have prevailed and would be totally appropriate in today's landscapes. What I really enjoy about the book, though, is the author's unapologetic and matteroffact acceptance of crabapples, warts and all. For example, he begins the chapter on insect and disease pests with this blunt statement: \"It should not be considered strange or disturbing that apples and crabapples are visited once in a while by some unwelcome guest.\" You have to admire Mr. den Boer's understated admission Crabapples 3 that certain members of the genus Malus do have pest issues, but the reality is few landscape plants are problemfree. Yet for some reason crabapples are subjected to much disrespect by certain detractors, even those who readily accept the premise that most landscape plants aren't perfect. crabapple nay sayers are happy to share their tales of crabapple woe, particularly when they involve suscepti bility to foliar diseases (\"My Uncle Vito over in Dubuque had a crabapple in his front yard that would defoliate completely every July.\") or fruit litter (\"You think that's bad ... my Aunt Betty had one that would drop loads of rotting, messy fruit all over her patio every summer.\") These repeated knocks against crabapples often trace to plantings of oncepopular, older crabapple cultivars such as `Hopa' and `Radiant'. Origi nally embraced for their headturning spring flower extravaganzas, these cultivars are now sadly, and maybe a bit unfairly, remembered only for debilitating disease problems and overly large, nonpersistent fruit. Unfortunately, a suf ficiently large population of `Hopa', `Radiant', and other lessthanstellar cultivars still can be found in present day landscapes, reinforcing the misperception that all crabapples defoliate in July and double as fastfood emporiums for every yellow jacket wasp in the neighborhood. But surely we--whether plant scientists or backyard gardeners--should understand the folly of making blanket statements about a group of plants with upwards of 900 named selections. After all, a family (in the non taxonomic sense) that large is bound to produce a few bad apples, if you'll excuse the pun. Why Crabapples Still Rule The fact is that crabapples remain atop the list of small ornamental trees used in residential and commercial landscapes in UsDA hardi ness zones 4 through 7 for many very good reasons. crabapples offer an avalanche of fra grant and colorful spring flowers in white and JeFF ILes Select apple-scab-resistant cultivars in order to avoid the heartbreak of mid summer crabapple defoliation. 4 Arnoldia 67\/2 2009 JeFF ILes Malus floribunda sports beautiful pink buds and white flowers. JeFF ILes Crabapples with persistent fruit provide months of color. Crabapples 5 NANcY ROse The weeping branches of `Red Jade' laden with bright red fruit. shades of red ranging from palest pink to deep burgundy. As an added spring attraction, many crabapples display beautifully contrasting col ors as the flower evolves from tight bud stage to fully opened flower--for example, deep pink buds opening to white flowers or deep red buds becoming bright pink flowers. Most crabapples have handsome foliage with leaf color ranging from dark green to burgundy. Though generally not noted for fall foliage color, some crabapples including M. tschonoskii and `satin cloud' develop eyecatching shades of orange, crimson, and purple, while others flaunt hues of apricot (`Prairie Maid') and goldenyellow (`Amberina' and `Red swan'). Providing as spectacular a display as their spring blossoms but much longer last ing, the best crabapples bear bushels of vividly colored fruit that enliven the fall and winter landscape. Another plus is the broad array of growth habits and mature sizes that makes it possible to choose a crabapple for practically any landscape situation. Finally, when planted on appropriate sites (welldrained soils and full sun) and given modest annual care, crabapples can have a functionally effective life of at least 40 to 50 years, and sometimes much longer. Where Do They All Come From? There are interesting stories behind the dis covery, naming, and introduction of every spe cies, variety, and cultivar of crabapple. From M. baccata, gleaned from the wilds of siberia and named by Linneaus in 1767, to modern cultivars that owe their existence to count less crosses and backcrosses, one has to marvel at the imagination, determination, and luck required to bring a single crabapple selection to the attention of the gardening public. As an illustration, consider the circuitous birthing path for the much admired weeping crabapple `Red Jade'. The `Red Jade' story begins in the early to mid 1800s in northeast Asia with the discovery and introduction of Malus prunifolia. The plumleaf crabapple was known for having many forms, and as luck would have it, a weeping form was 6 Arnoldia 67\/2 2009 JeFF ILes A Father Fiala introduction, `Orange Crush' crabapple is gaining popularity. discovered and given the cultivar name `Pen dula'. Later, M. prunifolia `Pendula' was crossed with M. floribunda (Japanese flowering crabap ple) with the result being a small, weeping tree eventually dubbed M. floribunda `exzellenz Thiel'. selected by spath Nursery in Germany and introduced to North America by the Arnold Arboretum in 1912, this diminutive, disease prone crabapple was one of the first weeping ornamental trees used in the United states. In 1935 serendipity stepped in as Dr. George M. Reed of the Brooklyn Botanic Garden either discovered or purposely germinated and grew openpollinated seedlings from M. floribunda `exzellenz Thiel'. What initially captured his attention isn't clear, but one of those seedlings developed into a beautiful weeping tree. In 1953 it was given the cultivar name `Red Jade'; the name remains a bit of a mystery but probably refers to the bright red, inch diameter fruit and the glossy \"jade\" green foliage, two notable and recognizable features of the cultivar. Now fastforward to one of today's rising stars, M. `Orange crush'. This delightful intro duction sports orangecrimson flowers, handfuls of deep maroon fruit, and excellent disease (and Japanese beetle) resistance. But its existence and subsequent rise to fame comes only after a mindnumbing series of crosses, ending finally when Father John Fiala crossed M. `Liset' with M. `Red swan'. And you can bet M. `Orange crush' will join the hybridization dance many times before it's put out to pasture. Selecting the Right Crabapple Finding a great crabapple for your landscape is pretty easy these days. The vast majority of crabapples now sold in nurseries and garden centers have much improved resistance to dis ease compared to their predecessors, and also feature highly ornamental fruit that is either small in size, persistent, or relished by our winged friends. The decision to include one or several crab apples in a landscape planting really hinges on several factors. First and foremost, the tree you choose must fit the site. For example, if you don't have sufficient room for a large tree (stan Crabapples 7 NANcY ROse Dense-crowned `Coralcole' (Coralburst) crabapple fits in smaller spaces. dard crabapples typically grow 20 to 25 feet tall and wide), you might consider one of several dwarf selections such as `camzam' (camelot), `cinzam' (cinderella ), `coralcole' (coral burst), or `Lanzam' (Lancelot). And if you like the somewhat formal look of dwarf forms top grafted to a standard, then you must investigate the aptlynamed `Lollizam' (Lollipop) and two Malus sargentii selections, `select A' (Firebird) and `Tina'. If you're looking for an uprightgrow ing selection that will pose minimal problems for pedestrian and vehicular traffic, the increas ingly popular `Adirondack' (selected by Don egolf at the United states National Arboretum) is the crabapple for you. But if space constraints aren't an issue (parks, golf courses, entryway plantings, large residential lots, etc.) imagine the visual impact of informally arranged drifts (5 to 9, or more) of redflowering `cardinal', red fruited `David', or goldfruited `schmidtcutleaf' (Golden Raindrops). Next, consider special maintenance issues such as disease susceptibility. In a perfect world, we'd quickly rule out using crabapple JeFF ILes White-flowered `Adirondack' has a tidy upright-vase shape. The crabapple to the right is `Purple Prince'. JeFF ILes NANcY ROse NANcY ROse Clockwise from upper left: `Camzam' (Camelot) `Jewelcole' (Red JewelTM) `David' `Donald Wyman' `Schmidtcutleaf' (Golden Raindrops) NANcY ROse NANcY ROse The \"Best\" Crabapples (Malus spp.) TAxA `Adirondack' FLOWeR white FRuiT orange-red (\") HT\/WD 18'\/10' FORm upright (inverted cone) FOLiAGe dark green Limitations: slow-growing `Camzam' (Camelot) fuchsia-pink burgundy (3\/8\") 10'\/8' rounded\/compact dark green\/burgundy Limitations: not much late-season interest `Cardinal' pinkish-red Limitations: none known `Cinzam' (Cinderella) white deep red (\") 16'\/22' broadly spreading dark purple-red gold (\") 8'\/5' rounded, upright light green Limitations: slow-growing `David' white scarlet (3\/8\") 15'\/20' rounded light green Limitations: alternate bloom; light apple scab noted; fruit mummies persist until spring `Donald Wyman' white bright red (3\/8\") 20'\/20' rounded medium green Limitations: apple scab noted; fruit mummies persist until spring floribunda pink-white amber (3\/8\") 12'\/20' spreading\/irregular medium green Limitations: unimpressive fall fruit display `Schmidtcutleaf' (Golden Raindrops) white golden-yellow (\") 20'\/15' upright medium green\/deeply cut Limitations: alternate-year bloom; fire blight has been reported `Lanzam' (Lancelot) white gold (3\/8\") 10'\/8' oval medium green Limitations: flowers\/fruit borne on interior of the tree which diminishes their ornamental effect `Louisa' rose-pink amber (3\/8\") 10'\/15' weeping dark green\/glossy Limitations: fruit are ornamentally insignificant `Orange Crush' rose-red red (3\/8\") 15'\/15' rounded purplish-green Limitations: none known `Prairie maid' deep pink red (3\/8\") 15'\/15' rounded medium green\/yellow-apricot fall Limitations: none known `Prairifire' pinkish-red dark red (\") 20'\/20' rounded purple turning reddish-green Limitations: requires pruning to correct overcrowded branching `Purple Prince' rose-red maroon (\") 20'\/20' rounded purple turning bronze-green Limitations: heavy fruit production may weigh branches down `Jewelcole' white (Red JewelTM) Limitations: none known `JFS-KW5' (Royal Raindrops) pinkish-red red (\") 15'\/12' pyramidal medium green red (\") 20'\/15' upright purple cutleaf\/orange-red in fall Limitations: none known 10 Arnoldia 67\/2 2009 for crabapple lovers because resistance to scab apparently is not a forever kind of thing (or, is not a permanent and bind ing contract between pathogen and host). In fact, all it takes is one lucky \"super\" ascospore infecting a previously resistant crabapple host to begin the pro cess of resistance breakdown in that host. Notable examples of resistance breakdown and the subsequent development of scab have occurred on Malus `Prairifire', `Bob White', `Jew elcole' (Red JewelTM), and floribunda. But sometimes positive attributes outweigh the nega tive, and therefore I'm willing This specimen of `Bob White' shows just a few spots of apple scab on its leaves. to look the other way when `Indian Magic' jettisons most of its scabflecked leaves in late summer, only to reveal one of the most visually stunning fruit displays in all of Malusdom (see front cover). When the topic of fruit size and persistence comes up, crab apple detractors fre quently trot out the poster child for obnoxious fruiting behavior, Malus `Dolgo' (ignor ing the fact that its large crim son fruits are great for making tasty preserves). But it would be disingenuous to paint all crabapples with the same brush. For example, crabapple selections like `Jewelcole' (Red if only Aunt Betty had planted her `Dolgo' crabapple (shown here) out in the yard JewelTM) and `Donald Wyman' instead of next to the patio she wouldn't have a crabapple mess underfoot. produce bright red, extremely selections with poor resistance to fungal patho persistent fruit that eventually fall from the gens Venturia inaequalis (apple scab) or Botrytree, but only after they've dried and shriv osphaeria obtusa (frogeye leaf spot) responsible eled to onehalf their original size. Others like for premature defoliation, and would never `snowdrift' and `Bob White' drop very little entertain the addition of a crabapple suscep fruit thanks to the work of opportunistic and tible to the bacterium Erwinia amylovora (fire grateful birds. And on those sites where fruit blight). The fungal prankster responsible for production of any kind is forbidden, fruit apple scab (actually, there are several races of less selections `spring snow' and newcomer V. inaequalis) has been especially frustrating `Jarmin' (Marilee) are viable options. NANcY ROse NANcY ROse Crabapples 11 The bright red fruit of `Donald Wyman' last through the winter. NANcY ROse The bite-sized orange fruit of `Snowdrift' attract birds. NANcY ROse 12 Arnoldia 67\/2 2009 NANcY ROse The form of weeping crabapple `Red Jade' outlined in snow. JeFF ILes The pinkish-red flowers of `Prairifire' crabapple. Crabapples 13 NANcY ROse What about weeping crabapples? Real or imagined, several barriers stand in the way of using weeping trees in the landscape. For start ers, consider the word \"weeping.\" Who wants a sad landscape? secondly, trees like weeping willow and weeping mulberry have, albeit unfairly, caused many weekend gardeners to be wary of any plant with cascading branches. Finally, and perhaps most importantly, weeping trees can be very difficult to integrate into the landscape. They vie for attention when used in groupings and look awkward and forlorn if used as a solitary specimen in the middle of a large lawn. And sticking one smack dab in the center of that unnaturallooking berm in your front yard isn't the answer either. But when classy, weeping crabapples like `Louisa' and `Huber' (Royal Fountain) come along, we are obliged to find them a prime location in the landscape where they can be viewed and appreciated at any time of day and throughout the year, espe cially during the winter months. Positioning a weeper at the corner of a home, near a water feature, atop a terrace, or at the end of a shrub border will gain approving looks from visitors and neighbors alike. Finally, consider a crabapple's ability to stop you in your tracks as you stroll through the landscape. If you've ever seen `Prairifire' awash in bright pinkishred flowers, `Doubloons' sport ing a bumper crop of goldenyellow fruit, the handsome purple cutleaf foliage of `JFsKW5' (Royal Raindrops), or the memorable silhou ette of weeping `Red Jade' encased in a glittering mantle of ice, well, you know what I mean. still, there are some who can't be convinced crabapples are anything but diseaseprone, messy trees. And in all honesty, this anti crabapple mindset is probably a good thing. I mean, what kind of crazy world would it be if everyone began planting crabapples? see you at the garden center. Resources Beckerman, J., J. chatfield, and e. Draper. 2009. A 33year evaluation of resistance and pathogenicity in the apple scabcrabapples pathosystem. HortScience 44(3): 599608. Brewer, J.e., L.P. Nichols, c.c. Powell, and e.M. smith. 1979. The Flowering Crabapple A Tree for All Seasons. coop. ext. serv. of Northeast states. Ne223, NcR 78. Delicate pink flowers flow along the weeping branches of `Louisa'. den Boer, A.F. 1959. Ornamental Crab Apples. The American Association of Nurserymen. Dayton, D.F. 1982. `Prairifire'. HortScience 17(2): 262. egolf, D.R. 1987. `Adirondack' crabapple. HortScience 22(5): 969970. Fiala, J.L. 1994. Flowering Crabapples: The Genus Malus. Timber Press, Portland, Oregon. Iles, J.K. and J.s. stookey. 1997. crabapples: sales trends and consumer preferences in Iowa. Journal of Arboriculture 23(3): 9499. Romer, J.P., J. K. Iles, and c.L. Haynes. 2003. selection preferences for crabapple cultivars and species. HortTechnology 13(3): 522526. Jeff Iles is Professor and chair of the Department of Horticulture at Iowa state University, Ames, Iowa. "},{"has_event_date":0,"type":"arnoldia","title":"Malus at the Arnold Arboretum: An Ongoing Legacy","article_sequence":3,"start_page":14,"end_page":21,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25470","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15eb36d.jpg","volume":67,"issue_number":2,"year":2009,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Malus at the Arnold Arboretum: An Ongoing Legacy Michael S. Dosmann NANCy RoSE Malus in bloom on Peters Hill, the Arnold Arboretum, May 2008. I n his book Flowering Crabapples: The Genus Malus, the late Father John Fiala (1994) states that \"no horticultural institution did as much for introducing and discovering new species, varieties, or special clones [of Malus] as did the Arnold Arboretum.\" Those are humbling words coming from such an authority as Father John. As I considered his accolade, I asked myself: What were the drivers that made this all possible? No doubt there were a number of factors involved in making the Arnold Arboretum \"the `mother arboretum' for flowering crabapples\" (Fiala 1994). Timing played a critical role in the initial development of the crabapple collection as well as its ongoing use and development. The Arboretum's founding in 1872 and early rapid expansion of collections coincided with vigorous plant exploration efforts around the world. With respect to Malus, countless taxa new to science were collected from the wild and described, while many others new to North America were introduced from cultivation else- where (primarily Europe). Additional introductions of taxa from varying parts of their native ranges ensured that a high degree of genetic variation was present. Simply having a diverse and sizeable collection of crabapples does not necessarily make it significant, however. The collection's active use in science throughout its existence put it on the map. Early on, the Malus collection was notably used in the study of taxonomy--the description of new species and their classification. This was followed by the collection's incorporation into better understanding genetics and cytology, as well as physiology. The collection proved to be of value to applied horticulture as well. Following World War II, as the demand for greater diversity of high-quality landscape plants increased, the products of these plant-breeding efforts (novel hybrids and cultivars) were grown and evaluated at the Arboretum. Development and scientific use of the collection was made possible by a number of prominent Arboretum personalities. Charles S. Malus at the Arnold Arboretum 15 ARNolD ARboRETuM ARChIvES Famed plant explorer Joseph F. Rock made this image of Malus transitoria on an expedition in Kansu (Gansu) province, China, on October 21, 1926. 16 Arnoldia 67\/2 2009 Sargent, first director of the Arboretum, knew the research value of a well-documented collection and ensured that the initial development of the Arboretum, including its growing repository of apples and crabapples, would get off on the right foot. he also recognized that Rosaceae was indeed too large a family to occupy its allotted space--the hillside currently known as State lab Slope near the Forest hills Gate--which was dictated by the Arboretum's design based on the bentham and hooker sequence of plant families. And so, at the end of the nineteenth century, he designated large expanses on Peters hill for the cultivation of Pyrus, his beloved Crataegus, and of course Malus. The expansion provided much relief, as numerous new species, hybrids, and cultivars were rapidly being introduced and needed space. Sargent himself collected and introduced new Malus, including the low-growing M. sargentii and the lesser-known but highly ornamental M. tschonoskii, both from his 1892 trip to Japan. Amazingly, the original specimens of these two species, now nearly 120 years old, still grow near ARNolD ARboRETuM ARChIvES the bradley Rosaceous Collection and represent the Arboretum's oldest Malus accessions. Ernest h. Wilson also played the role of explorer and introducer. Plantae Wilsonianae credits Wilson with collecting from some 16 Malus species during his travels in China, several of which were taxa new to science. Perhaps the best of these is Malus hupehensis, the picturesque small tree with a vase-shaped habit that Wilson made numerous collections of during both his veitch and Arboretum expeditions. In describing its merits, Wilson (in Sargent 19131917) notes that \"it is very beautiful in spring when covered with light pink flowers, and resembles at this time a flowering cherry rather than an apple tree; the effect of the flowers is heightened by the purple calyx and the purplish tints of the unfolding leaves.\" Alfred Rehder, Arboretum taxonomist, may not have collected and introduced material from the wild, but he certainly applied his shrewd skills of observation and classification in describing and naming scores of the new Malus Malus floribunda on Peters Hill, photo by Ralph W. Curtis, May 10, 1922. Malus at the Arnold Arboretum 17 MIChAEl DoSMANN MIChAEl DoSMANN The lovely pink flowers of Malus hupehensis. One of the original Malus `Mary Potter' (181-52-B), planted in 1952. ARNolD ARboRETuM ARChIvES species and countless infraspecific taxa and hybrids. hybrids within Malus are quite common, and as the Arboretum's collection grew and diversified, genes began to mix, hybrids arose, and more discoveries were made. Perhaps the most ardent scientific user of this botanical petri dish was Karl Sax, former Arboretum director and research scientist at the bussey Institute. Through the course of much of his Arboretum career, he integrated the Arboretum's Malus collection into a wide array of studies ranging from polyploidy and apomixis (Sax 1959) to plant physiology (Sax 1957). A byproduct of his many cytology and breeding experiments was an abundance of hybrids, from which Sax was able to evaluate and select a number of crabapple cultivars (Sax 1955). Four prominent In this 1959 photo by Heman Howard, Karl Sax is seen with a grafted dwarf apple ones are `blanche Ames', `hen- tree, one of his many research interests at the Arboretum. rietta Crosby', `henry F. du Pont', and `Mary Potter'. The latter is perhaps of single white flowers in the spring and bright his finest introduction and a personal favorite red fruit in the autumn. Making the story all of mine. `Mary Potter'--a cross between M. sarthe more interesting is that it was named after gentii `Rosea' and M. x atrosanguinea--is lowthe daughter of C. S. Sargent, and has the Sargrowing yet spreading, producing an abundance gent crabapple as a parent. 18 Arnoldia 67\/2 2009 ld, robust collections like the Arnold's are always full of new surprises. An interesting story concerns two unusual trees growing on Peters hill, AA 691-52-A and b. While a Putnam Fellow in the spring of 2001, I became enamored by their wide-spreading, low-branching form; 691-52-b, the slightly larger of the two, stands 18 feet (5.5 meters) tall and 33 feet (10.1 meters) wide. The leaves and flowers are borne in dense, tight clusters throughout the canopy, giving the two specimens an unusual cloudlike appearance. The flower buds are magenta at first, and then transition into light pink before they open into creamy white blooms. The tag read simply \"Malus sp.\" so I figured the trail was cold and that nothing more could be found about these plants. however, hidden away in the records was the note: Sax 7841. \"Sax Numbers,\" as these were known, were remnants of Karl Sax's own accessioning system at the bussey Institute and referenced his research plants or crosses (this one being the 78th plant or cross of 1941). but unfortunately, no additional documentation had ever been found that explained the numbers further, such as source of material, what the parentage had been if it was a cross, or what the understock or scions may have been in one of his experiments. Another seeming dead end, I gave up on pursuit of this additional information. Nearly a year later, though, while rummaging through the archives, I stumbled upon an unknown notebook of Sax's that turned out to be his master list of hybrids and experimental units. With this fortunate The mystery crabapple: Malus 691-52-A. find, I was able to identify not only these two plants but also a great number of other hybrid Malus, Forsythia, Prunus and other genera. It turned out that the duo in question were hybrids that Sax had made between M. lancifolia and M. sylvestris. Although I do not know if it was his original intent when making the cross, he used these hybrids in a rootstock experiment, possibly to examine any potential dwarfing effects rootstocks can have upon the scion above. Two seedlings of Sax 7841 were the ungrafted individuals I was struck by (691-52-A and b), while 780-52-A and D, located westward and up the hill a bit, were grafted plants that had Sax 7841 as the understock and an unknown wild apple as the scion (his notebook did not provide that detail, alas). Although Sax's cross yielded an unusual plant with ornamental habit, it would be premature to introduce it as a cultivar without further evaluation. And so, in 2007, Arboretum propagator Jack Alexander grafted budwood from both plants of 691-52 onto numerous seedlings of Malus `Antonovka'. Soon these trees will be planted and further evaluated for potential selection and introduction. o A Malus Mystery NANCy RoSE Malus at the Arnold Arboretum 19 While Sax may have been the creator of many of the cultivars, it was Arboretum horticulturist Donald Wyman who was their biggest promoter. he lauded their merits throughout the pages of Arnoldia and in his books, and advocated for their use in his lectures and correspondence. And, like Sargent before him, Wyman tapped his extensive global horticultural network to distribute Arboretum selections as well as acquire new taxa to grow and evaluate. In honor of Wyman's dedication to crabapples, the Arboretum introduced Malus `Donald Wyman' in 1970 to honor him in his retirement. A fantastic selection, it is appreciated for its abundant white flowers in the spring, relatively high disease resistance, and very long-lasting display of brilliant red fruit from autumn through winter. Interestingly, this tree was actually a spontaneous seedling that was first recorded growing on Peters hill on March 20, 1950. Due to its aesthetic appeal, it was later accessioned and then selected and introduced as the cultivar known today; the original tree still stands. It is ironic that, despite the great efforts of breeding and The original specimen of `Donald Wyman' (seen here in spring bloom and fall fruit) selection made over the years, still stands on Peters Hill. the Arboretum's most important crabapple introduction to date must be For example, the 1980 Sino-American botanical chalked up purely to serendipity. Expedition yielded several fascinating collecAlthough the period from Sargent to Wyman tions, including an unusual southern provemay have been known as the \"Golden Era for nance of M. baccata, the Siberian crab, found Crabapples\" at the Arboretum, work in the in hubei province. In addition to its unusual collection did not end when Wyman retired. collection site, this collection (SAbE #1298) As the Arboretum shifted the focus of its colproduces flowers and fruits borne on particulections policy towards acquisitions of known larly long pedicels (Spongberg 1991). An amazwild origin in the 1970s and 1980s, novel germing trio of this accession, AA 1843-80-D, h, and plasm from Asia again crossed the threshold. I, each with outstanding spiral-grained bark, MIChAEl DoSMANN MIChAEl DoSMANN 20 Arnoldia 67\/2 2009 Resistance can be conferred by the presence of the Vf gene, whose original source came from Malus floribunda selection 821 growing at the university of Illinois. This clone, the most frequently used source for scab resistance in the world (Koller et al. 1994), arose from seed sent from the Arboretum in 1908 to C. S. Crandall, a geneticist at the university of Illinois who was studying inheritance patterns in Malus. however, it was not until the 1940s that the initial crosses were evaluated for disease resistance, and it has only been in the last 30 years that high-yielding cultivars have been introduced through the PRI (Purdue-Rutgers-Illinois) Apple breeding Program, the most important just in the last few years (Janick 2006). I like this story for a number of reasons. It demonstrates how important it is for the Arboretum to distribute material (plants, seeds, cuttings, tissue, etc.) to researchers to enable their work. It also illustrates the importance of prudence and patience when working with trees--in this case, it has taken nearly 100 years since the original shipment from the Arboretum for the most meaningful dividends in research (in this case superior apple cultivars through one breeding program) to be realized. Currently, the Arboretum's living collection of Malus comprises 455 accesThis trio of Malus baccata display their distinctive spiral-grained bark sioned plants (about 3% of the total in the Arboretum's Bradley Rosaceous Collection. collection), representing 173 unique taxa, can be found in the bradley Rosaceous Collec104 of which are cultivars. Development is contion. other collections of M. hupehensis and stant: old lineages of high value are maintained M. halliana were made on this momentous through vegetative propagation, discretionary expedition as well, significantly increasing the accessions are disposed of, and new germplasm genetic diversity of these species in cultivation. is obtained. Recent and future renovations on The Arboretum collection continues to undergo Peters hill and the bradley Rosaceous Collecdevelopment. Recently, we have acquired a tion provide wonderful opportunities to grow number of wild-collected M. sieversii, the pronovel material of both wild and cultivated origenitor of the cultivated apple found growing in gin. At the species level, the goal is to possess Kazakhstan and neighboring countries. two to three wild provenances; for cultivars, With respect to the enhancement of fruiting we will continue to trial new introductions of genotypes, the Arboretum's collection played a ornamental selections and will also begin to noteworthy role, even if it was indirect. Apple feature several selections of eating apples. And, scab is a serious fungal disease that damages of course, the collection will continue to hold not just the leaves of trees but also fruits, causmany old and historically important selections, ing serious economic losses in apple orchards. including those introduced by the Arboretum. MIChAEl DoSMANN Malus at the Arnold Arboretum 21 Crabapple Cultivars Introduced by the Arnold Arboretum `Barbara Ann' `Dorothea' `Henrietta Crosby' `Henry F. Dupont' `Katherine' `Pink Pearl' `Blanche Ames' `Bob White' `Donald Wyman' `Mary Potter' `Prince Georges' M. baccata `Columnaris' M. baccata `Jackii' M. ioensis `Palmeri' M. x robusta `Erecta' M. sargentii `Rosea' M. x zumi `Calocarpa' Malus `Dorothea'. NANCy RoSE Literature Cited: Fiala, J. l. 1994. Flowering Crabapples: The genus Malus. Timber Press, Portland, oregon. Janick, J. 2006. The PRI apple breeding program. HortScience 41(1): 810. Koller, b., l. Gianfranceschi, N. Seglias, J. McDermott and C. Gessler. 1994. DNA markers linked to Malus floribunda 821 scab resistance. Plant Molecular Biology 26(2): 597602. Sargent, C. S. 19131917. Plantae Wilsonianae: An enumeration of the woody plants collected in western China for the Arnold Arboretum of Harvard University during the years 1907, 1908, and 1910 by E. H. Wilson. Cambridge university Press, Cambridge (united Kingdom). Sax, K. 1955. Plant breeding at the Arnold Arboretum. Arnoldia 15(2): 512. Sax, K. 1957. The control of vegetative growth and the induction of early fruiting of apple trees. Proceedings of the American Society for Horticultural Science 69: 6874. Sax, K. 1959. The cytogenetics of facultative apomixis in Malus species. Journal of the Arnold Arboretum 40: 289297. Spongberg, S. A. 1991. A Sino-American sampler. Arnoldia 51(1): 214. Michael S. Dosmann is Curator of living Collections at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"In the Footsteps of Father David","article_sequence":4,"start_page":22,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25469","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15eb328.jpg","volume":67,"issue_number":2,"year":2009,"series":null,"season":null,"authors":"Basset, Cedric","article_content":"In the Footsteps of Father David Cedric Basset A rmand David (18261900), famously known as \"Father David,\" is well known by those passionate about plants. Indeed, many plants carry his name, such as Davidia and Acer davidii. Though best known for his plant discoveries, one cannot mention this great figure without also mentioning the famous giant panda that he discovered in 1869 near Baoxing (previously Moupin) in the Sichuan province of China. During our expedition to Sichuan in May, 2007, we followed the same paths that Father Armand David took during the second half of the nineteenth century. These regions--with their extraordinarily rich flora and fauna--are fortunately still preserved, no doubt in part because they remain very difficult to access. A Few Notes from Father David Upon arriving in Moupin, Armand David wrote: \"The land of steep mountains is, despite the loggers and farmers, abundantly forested with fir trees and cedars up to 3,000 m ... The lanceolate pine and the narrow-leaved pine, as well as the alder of Setchuan, thrive up to 2,000 m. The rhododendrons are particularly abundant.\" It All phoTogrAphS Are By The AUThor This flowering Rhododendron was part of the extreme botanical richness we admired in the narrow valley of Pujigou, located south of the nature reserve of Fengtong. Armand David 23 should be noted that there are no cedars (Cedrus) in this region; Father David probably uses this term to designate other conifers with a similar horizontally spreading form. The lanceolate pine is certainly his designation for Cunninghamia lanceolata. At that time, Moupin (now known as Baoxing) was still part of Tibet. And Armand David wrote: \"Long closed to the Chinese, the principality now tolerates their growing number.\" That was a different era, indeed. on March 23, 1869, having just discovered the giant panda (Ailuropoda melanoleuca) he writes: \"The young bear is entirely white, except for his four limbs, his ears, and the area around his eyes which are a deep black. Thus, we have here a new species of Ursidae that is very remarkable not only because of its color but also because of the hairiness under its paws.\" later, he writes concerning his botanizing: \"The large rhododendrons are flowering, and I can already distinguish at least seven distinct species. I also found, in the middle of a wet forest, a magnificent magnolia with large purplish flowers and with no leaves yet.\" This may be Magnolia liliiflora, naturally present in this region. Baoxing, Town of the Panda Nowadays, the little town of Baoxing Cunninghamia lanceolata is a large conifer in the cypress family (Cuprespays homage to Father David with a saceae) that can reach 50 meters (164 feet) in height. It is present in the statue of him and with another that landscape of the Chinese provinces explored by Father Armand David. celebrates his discovery of the panda. The balustrades along the river are engraved Wolong, pass to the south and north of Baoxing, with representations of the numerous species respectively, while the northsouth road that of plants and animals that he discovered during connects rilong to ya'an through Baoxing is his sojourn in the region. poorly travelled. According to numerous local officials, There are several explanations for this lack Armand David's discovery over 130 years ago of tourist traffic. The road linking rilong to confers on Baoxing the status of \"cradle of the ya'an is not always in a good state. It is long giant panda.\" And yet, Baoxing remains infreand winding, and the lack of bus service forces quently visited by tourists. Westerners are rare, one to use a taxi. There are few possible stops since the town is located on a road little used along the road. our stop in the small town of by tourists. The roads that connect Chengdu to yanjingping was an adventure: no real hotel, Tibet through Kangding and litang, or through only one very dirty house, and one building 24 Arnoldia 67\/2 2009 ArNolD ArBoreTUM ArChIveS An Extraordinary Discoverer of Life rmand David was born September 7, 1826, in the village of espelette in southwestern France. on November 4, 1848, he joined the lazarist order in paris where he studied for several years. he then traveled to Italy to study medicine, zoology, and botany. on July 5, 1862, he arrived in peking where he lived for the following twelve years. During those years, he carried out three expeditions to western China. After falling sick during the third expedition, he returned to France in 1874. During his life in China, he visited Inner Mongolia, Shanghai, the Sichuan provinces, and hubei and Jiangxi, combining his missionary work with his scientific research. From March 1, 1869, until 1872, he worked in Moupin (now known as Baoxing) in Sichuan. During his travels in China, Armand David collected 13,000 specimens including 189 new plant and animal species, among these the handkerchief (or dove) tree (Davidia involucrata), the butterfly bush (Buddleja davidii), Lilium davidii, Populus davidiana, as well as thirteen species of rhododendrons, three magnolias, four firs, and four oaks. A where we found a room with no bathroom facilities. In the only restaurant in town, we involuntarily attracted a crowd and became, for the duration of our dinner, the main attraction. Baoxing, on the contrary, turned out to be a quiet small town, ideal for an enjoyable stop. There we stayed in a comfortable hotel where the rooms were very clean. The Forgotten Valley and Pujigou Baoxing is located to the south of the nature reserve of Fengtong. This reserve covers 40,000 hectares (98,842 acres), with 13 percent of the area serving as habitat for the panda. The town proudly advertises the region and its natural marvels--virgin forests, waterfalls, forests of Osmanthus, panoramas--but public transportation, as often in China, is non-existent. one must hire a vehicle and driver (fairly easily done at the train station) and communicate to the driver that he must wait all day or return to a meeting point after several days. otherwise, in the small villages, one would not be sure of finding a vehicle available for returning to town. We decided to enter the nature reserve by an alternative route, by taking the road that leaves from the northwest of Baoxing and winds to its final destination, the village of pujigou. The road, paved at its start, rapidly gave way to a narrow dirt path where we had perilous crossings with the trucks from a nearby quarry. To our surprise, after two and a half hours of driving, our driver stopped before a dilapidated wooden bridge and told us that we had to continue on foot. he told us that pujigou was located about an hour's walk further. We took five hours, since we walked very slowly at first, our botanical passion ignited as we marveled at discovering an interesting plant with each step. The flora in this infrequently traveled area offers a rare diversity, the very acidic soil being favorable to the growth of many plants of the ericaceae and the climate allowing amazing sub-tropical species to flourish. Armand David 25 We arrived at an old, abandoned building in the middle of the forest where two men and one woman lived without electricity. The reception was icy at this abandoned and empty inn. We were in pujigou. It is not really a village, but rather the remains of what must have previously been a remote mountain refuge. Deciding to flee this place, we turned around and went on to find a village where we were hosted by a local resident. The Fengtong Reserve In all of our previous trips to China we had never found such a wild valley as at the Fengtong reserve. Unlike more accessible nature reserves such as Wolong, here at Fengtong there was no road, no cars or buses, only a small path. The valley is narrow, with steep slopes covered with dense vegetation that benefits from the very humid air. From a botanical perspective, it is a real treat. The orchid Calanthe tricarinata grows about 30 centimeters (about 12 inches) tall. It enjoys semi-shaded areas and a humid climate. The superb striped bells of Enkianthus deflexus. Davidia involucrata (center) bloomed among the dense vegetation in this narrow valley in the Fengtong Reserve. All along the trail, magnificent handkerchief trees (the famous Davidia involucrata, dedicated to Father David) in full bloom hung over us. The edge of the path was full of flowering Disporum bodinieri (a member of Convallariaceae) and a somewhat rare Paris, Paris fargesii. In the nooks of dead tree trunks and on rocks, beautiful orchids--Calanthe tricarinata and Pleione limprichtii--bloomed abundantly. Above our heads we saw two beautiful shrubs, Dipelta yunnanensis of the honeysuckle family (Caprifoliaceae) and Enkianthus deflexus of the family of the rhododendrons (ericaceae). The giant dogwoods (Cornus controversa) spread their tiered silhouettes above the shrubs. 26 Arnoldia 67\/2 2009 the viburnum of China, Viburnum chingii. We met more frequently another little shrub with lots of flowers: Deutzia glomeruliflora. In this gorgeous reserve, another seasonal spectacle was provided by climbing plants of the lardizabalaceae: Holboellia and Akebia. Certain stems, several meters tall, were covered with flowers exuding a sublime scent. A few plants of Akebia trifoliata revealed flowers that were almost black. A little higher, Sinofranchetia chinensis, belonging to the same family, was reaching even farther up into the trees. The trail, although inaccessible to cars, was very good for walking. Certain signs showed that it was previously accessible to vehicles. The reserve is home to the giant pandas, and large stands of bamboo of the genus Drepanostachyum bordered the trail. We also saw a beautiful, large Yushania on which climbed Codonopsis tangshen (in Campanulaceae), not in flower. Cornus controversa displays its elegant horizontal branching habit. Some Rare Finds one great surprise was finding dozens of plants of one of the most spectacular hornbeams, Carpinus fangiana. I had wanted to see it for a long time and had already searched for it, notably at Mount emei (emei Shan). This tree is surprising for its large leaves (longer than 20 centimeters [7.9 inches]) and catkins that can reach 50 centimeters (19.7 inches) long. Several species of viburnum (among these Viburnum brevitubum) carried their long, white tubular inflorescences in the manner of Remembering Father David At the forest's edge and along paths on shady rocks, several species reminded us of Father David: Epimedium davidii, a small epimedium (Berberidaceae) with beautiful four-pronged yellow flowers. Acer davidii, David's maple (Sapindaceae), with its bark finely striped with white. Corydalis davidii (Fumariaceae) with its pretty yellow flowers. Much rarer is the impressive Corydalis anthriscifolia, a large plant with long purple inflorescences of which we saw only one specimen. In another small, narrow valley, we observed large arisaemas in flower with enormous leaves Armand David 27 The superb Carpinus fangiana growing in the dense forests of Pujigou. Holboellia sp., a perennial vine with strongly perfumed flowers. Viburnum brevitubum growing in the cracks of rocks in the Fengtong Reserve. composed of three leaflets. This was Arisaema dilatatum, a little-known species distributed from western Sichuan to Bhutan. We also were very surprised by the diversity of maple species in this valley. We admired the very rare Acer sutchuenense, a small tree of 5 to 8 meters (16.4 to 26.2 feet) high with trifoliate, denticulate leaves. The Big Surprise The exploration of this fantastic valley ended in an exciting discovery--finding plants of the famous and rare hellebore of Tibet (Helleborus thibetanus). This species (in ranunculaceae) lives in isolation, as the next closest hellebore species grow more than 5,000 kilometers (3,100 miles) to the west. This hellebore is doubtless the most delicate of its type, with sepals much Yet another plant named in honor of Father David, the lovely Epimedium davidii. more finely-textured than the other species. Its flowers vary from pinkish-white to dark pink, often with darker pink veins. Helleborus thibetanus was described by Franchet in 1885 from specimens collected in 28 Arnoldia 67\/2 2009 1869 by Armand David at Baoxing in Sichuan. The same year, Beresowski collected specimens in the province of gansu that were described by Maximowicz in 1890 under the name Helleborus chinensis, a name now synonymized under H. thibetanus. Tibetan hellebore's introduction to europe is relatively recent. In 1991, seeds were sent from China by professor Kao pao-chung of the Chengdu Institute of Botany. They had been collected from Sichuan, near Baoxing, on Dengchigow mountain at an elevation of 2,300 meters (7,500 feet). Among the long list of other species that we observed, we should mention Cotoneaster moupinensis, several superb dark Cardamine along the creeks, several Clematis, several Euonymus, a beautiful Magnolia that was not The rare Helleborus thibetanus bears delicately veined pink flowers. flowering, several honeysuckles (Lonicera), one Sorbus with simple leaves, and several plums (Prunus spp.). A Good Inspiration Father David chose well in coming to this mountainous region. The diversity of its flora and fauna is fascinating and important. efforts to protect the panda and the creation of reserves have allowed the preservation of very speciesrich valleys. To travel in this area, one must temporarily do without some comforts, but it is truly worth it, especially since Baoxing has several good hotels. This region is representative of western Sichuan on the road to Tibet. In addition to Baoxing, one should stop in Kangding, ganjia Caoyuan, garze, or also at Mugue lake to discover fabulous landscapes and a very diverse natural world. Bibliography Boutan, e. 1993. Le nuage et la vitrine: Une vie de Monsieur David. editions Chabaud, Bayonne. Wu, Z. and p. raven, eds. 1994+. Flora of China. Missouri Botanical garden press, St. louis. Cedric Basset is responsible for the collections of the Botanical garden of lyon in lyon, France, and is a specialist in Asian flora (www.asianflora.com). Translated from French by elizabeth h. Zacharias, ph.D., and Ian C. Bourg, ph.D. The very strange-looking Arisaema dilatatum. "},{"has_event_date":0,"type":"arnoldia","title":"Book Excerpt: Between Earth and Sky: Our Intimate Connections to Trees","article_sequence":5,"start_page":29,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25467","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15eab6b.jpg","volume":67,"issue_number":2,"year":2009,"series":null,"season":null,"authors":"Nadkarni, Nalini M.","article_content":"Book excerpt: Between Earth and Sky: Our Intimate Connections to Trees Nalini M. Nadkarni. University of California Press, Berkeley and Los Angles, California. 2008. 322 pages. ISBN 978-0-520-24856-4 Editor's Note: Dr. Nalini Nadkarni is a noted expert on the ecology of forest canopies in both tropical and temperate regions. Her innovative research has led to greater understanding of forest canopy ecology and forest ecosystem ecology. In this book, Nadkarni steps back from pure science and instead explores the profound connections between humans and trees. The book's structure is based on a creative modification of psychologist Abraham Maslow's pyramidal hierarchy of human needs. Here, the pyramid represents levels showing how human needs are met by trees, from the basic levels of physical needs like food and shelter, mid levels including imagination, language, and connections to time, and ultimately to the apical levels of spirituality and mindfulness. The following are brief excerpts from three chapters. Goods and services lthough humans now rely on ships and airplanes made of metal for long-distance transport, trees still figure into the regional and local transportation of our commodities. In 2000, for example, well over half of the $1.7 trillion worth of goods that entered and left the United States used some form of solid-wood packing material, such as pallets and crates. In 2001, an estimated two billion pallets were in use in the United States--six for every American. Over half of these are designed to make just one trip, and pallets as a whole average just 1.7 trips. Only about 10 percent are recycled, ground up and used as landscaping mulch, animal bedding, or core material for particle board. The wood in the pallets that are discarded each year is enough to frame 300,000 average-sized houses. Each year, too, 500 million more pallets are made, consuming trees on the equivalent of 18,000 acres. Our global reliance on pallets also introduces nonnative pests. One is the Asian longhorned beetle, an \"exotic\" pest that has threatened North American hardwood trees such as maple, elm, birch, poplar, and willow since 1996. The clue that these large beetles arrived in \"Trojan pallets\" was that outbreaks were concentrated near warehouses in New York, New Jersey, and Chicago, which contain pallets from China and Korea, where the beetles are native. Since then, infested pallets have been intercepted by vigilant entomologists in many North American cities, and so far, serious outbreaks have been contained. Europe, meanwhile, is suffering from an invasion of the pinewood nematode, thanks to products received from the United States, China, and Japan. Because of such threats, many export companies have begun to use metal or plastic pallets. These in turn create other problems, as those materials are not as easily recycled. A 30 Arnoldia 67\/2 2009 T play and imaGination dreaminG aloft o many climbers, the ultimate experience is spending a night high in the treetops. There is something about sleeping in the forest--whether on the ground or in the trees--that brings us as close as we can get to nature. My first overnight experience suspended in a hammock between branches of a giant tropical rainforest tree remains vivid in my memory even thirty years later. I climbed into the canopy as the sun set, the darkening understory giving way to the lighter environment of the canopy--though that, too, gradually became part of the jungle night. Bird songs gradually yielded to the buzzing, whirring, creaking calls of insects, which grew louder both below and above me, a sort of stereo effect I had never heard before. I curled up on my hanging cot, water bottle and a bag of snacks tied to an auxiliary cord, my harness and rope giving me a sense of security as the spookiness of being two hundred feet above the ground crept into me. At some point during the night, an anteater rambled over to my perch, in pursuit not of a dormant human but rather of the steady stream of leaf-cutter ants that were harvesting chunks of leaves from the trees and walking them along the branch highways down to their underground nests. On seeing me, the colliesized mammal seemed as startled as I had been. But we looked at each other for a long moment without fear, two arboreal animals in a high place on a dark night. Since that time, I have spent many nights aloft. What has surprised me is not the \"otherness\" of the canopy night compared to where we ground-bound humans normally sleep, but rather how homey and comfortable it seems up there with darkness stretching out in three dimensions. We were raised with the classic lullaby, \"Rock-a-bye baby in the treetops,\" with its inevitable and sobering conclusion: \"and down will come baby, cradle and all.\" And there are noxious insects and poisonous reptiles somewhere up there. But during those nights I spent on my canopy cot, swaying slightly in the wind one hundred feet above the ground, I couldn't have felt safer and more ready for sleep, lulled by my nocturnal companions above, below, and around me. connections to time rees express time with a precision and beauty that are unmatched in nature. Changes in their foliage mark the passage of Earth's seasons, while the incremental growth in their rings mark Earth's years. Nothing more effectively indicates seasonal transitions than the tender green of the emerging buds of spring, the rich, deep greens of summer, the multicolored leaves of autumn, or the delicate filigree of snow on tiny twigs after a winter storm. We are inspired by trees' relationships to time: the great age they can attain and the fierce disturbances they can endure. When we walk along the winding paths of a cemetery, we pass beneath the trees that have dwelt there far longer than their interred neighbors, giving us a comforting sense of continuity. Measuring the age of trees in tropical forests has been a perennial problem for ecologists. Because in many places favorable growth conditions occur year-round, many trees in the tropics are constantly growing and so have no rings at all. Even trees that do undergo seasonal growth, such as those in habitats where rainfall is concentrated into a few months of the year, have unreliable rings because they can jump into a growth mode in response to even small inputs of out-of-season rainfall. The tropical dendro- T Book Excerpt 31 chronologist, therefore, must turn to other methods to determine a tree's age. One surprisingly useful tool for tree dating emerged through the development of atomic weaponry. During the early era of nuclear testing, atomic devices were detonated in the atmosphere. The radioisotopes ejected from these explosions spread worldwide, forming a thin, weakly radioactive blanket over the earth. Some of these radioisotopes mimic naturally occurring elements so closely that many plants and animals cannot distinguish them. Trees take them up and incorporate them into their cells, along with their regular nutrients. Radioactive strontium, for example, mimics calcium, a nutrient that plants use to build new cell walls, much as animals use calcium to build bones. In 1954, trace amounts of radioactive strontium generated from bomb tests wafted through the air, dissolved in rain, entered the water cycle, were absorbed by roots, and then were incorporated into the living tissues of trees. This resulted in a short-lived but distinctive radioactive signal that has been held in the tissues of all of the trees living in the world at that time. Now, half a century later, scientists extract pieces of wood from tropical trees and note where in the cross-section of the trunk the \"1954 bookmark\" of radiation occurred. This allows scientists to measure how much each tree has grown since that time. Although the results cannot be extrapolated to determine how old a tree is, they do provide the dendrochronologist with an exciting tool to compare the rates of growth (from 1954 to the present) of individual trees and different species of trees that lack reliable rings. By revealing relative growth rates, this approach gives scientists a better understanding of population dynamics within forests. Adapted from Between Earth and Sky: Our Intimate Connections to Trees, by Nalini Nadkarni, published by the University of California Press. 2008 by the Regents of the University of California. "},{"has_event_date":0,"type":"arnoldia","title":"Autumn's Harbinger: Acer rubrum 'Schlesingeri'","article_sequence":6,"start_page":32,"end_page":32,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25466","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15eab26.jpg","volume":67,"issue_number":2,"year":2009,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Autumn's Harbinger: Acer rubrum `Schlesingeri' Michael S. Dosmann A utumn is my favorite time of year, and during the dog days of late summer I particularly look forward to the cooler, crisper, colorful months to come. That's why I am delighted when, on some sultry August afternoon, I notice that our Acer rubrum `Schlesingeri' has begun to express the first hints of leaf color at the Arboretum. In most years, the green foliage of this early-coloring red maple shades to bronze by mid August, and by early September the entire canopy is ablaze in carmine red. The colorful display usually holds into October. The precocious and stunning autumn coloration of this selection first caught the eye of Arboretum director Charles S. Sargent in the late 1800s. The original tree grew at the home of Sargent's neighbor, Mr. Barthold Schlesinger, in Brookline, Massachusetts. On February 13, 1888, budwood from this tree arrived at the Arboretum and, upon grafting, became accession 3256-A. It was planted along Meadow Road across from the Hunnewell Building, where it remains to this day. Curiously, this cultivar's introduction to the ornamental scene occurred not in North America but in Europe. Sargent had shared it with the world-famous Spath Nursery in Berlin, which first made it commercially available in their 18961897 catalog. During World War II, the nursery dissolved, no doubt limiting the supply of this sought-after clone. In 1951, the Arboretum distributed plants to some 25 cooperating nurseries as a means of promoting the cultivar and increasing supply. In his description of the tree and this distribution program, Donald Wyman (1956) noted the efforts made to learn if the precocious fall color trait was truly genotypic or just a function of environment: \"... scions from this variety were grafted on seedling red maples, but both the scion and the understock were allowed to grow. In the fall, it was clearly evident that the variety schlesingeri [`Schlesingeri'] would produce autumn color several weeks before the seedling understock on which it was growing, regardless of where it was planted.\" Unfortunately this cultivar is now often misidentified, so the Acer rubrum `Schlesingeri' that you purchase at the local nursery may not be true-to-type. This has even happened at the Arboretum. In the early 1980s, three trees labeled as `Schlesingeri' were donated by a large, reputable, national nursery. But in 1989, Arboretum horticulturist Gary Koller noted that they \"do not match 3256-A ... identification (of) this cultivar is questionable.\" Further observations proved Koller correct and these trees were duly removed. Michael Dirr, in his Manual of Woody Landscape Plants, also noted that \"some of the material in today's market does not appear similar to the Arnold Arboretum's fine specimen.\" And an interesting study on red maple cultivar coloration (Sibley et al. 1995) yields further evidence: although the trees of `Schlesingeri' examined in the study were obtained from a reputable nursery, they developed the wrong leaf color (orange) far too late (no earlier than the 5th of October) to be true `Schlesingeri'. Over 120 years later, this old sentry remains in its original location. It stands 65 feet (19.8 meters) tall with a crown spread of about 60 feet (18.3 meters), and its trunk diameter (below the lowest branch) is 44.6 inches (113.3 centimeters). Red maples generally reach maturity at around 75 years of age, so it is no surprise that this individual is in decline. Recent efforts to maintain this important lineage by rooting cuttings have been a success: accession 408-91-A grows next to Faxon Pond, and scores of new cuttings are now rooting in the greenhouse. One of these new plants will eventually replace the original tree, while others will be distributed to commercial nurseries so that they, too, will have the real cultivar again. References Sibley, J. L., D. J. Eakes, C. H. Gilliam, G. J. Keever, and W. A. Dozier Jr. 1995. Growth and fall color of red maple selections in the Southeastern United States. Journal of Environmental Horticulture 13(1): 5153. Wyman, D. 1956. New and rare ornamental plants recently distributed by the Arnold Arboretum. Arnoldia 16(79): 3351. Michael S. Dosmann is Curator of Living Collections at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23415","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160a76e.jpg","title":"2009-67-2","volume":67,"issue_number":2,"year":2009,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Cydonia oblonga: The Unappreciated Quince","article_sequence":1,"start_page":2,"end_page":9,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25461","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14e8926.jpg","volume":67,"issue_number":1,"year":2009,"series":null,"season":null,"authors":"Postman, Joseph","article_content":"Cydonia oblonga: The Unappreciated Quince Joseph Postman The quince of Persia attains a weight of 1.5 kilos (more than 3 pounds), ripens on the tree or in the store, and can be eaten like a soft ripe pear, according to a report in The Horticulturist, and Journal of Rural Art and Rural Taste of 1849 (Meech 1908). \"the quince, the `Golden Apple' of the ancients, once dedicated to deities and looked upon as the emblem of love and happiness, for centuries the favorite pome, is now neglected and the least esteemed of commonly cultivated tree fruits.\" (Hedrick 1922) Luther Burbank took credit for transforming this neglected fruit from a commodity that was \"altogether inedible before cooking\" into a crop he likened to the best apple. He half-jokingly cited a formula to make quince fruits edible prior to his breeding efforts: \"Take one quince, one barrel of sugar, and sufficient water...\" (Whitson et al. 1914). Burbank released several improved cultivars in the 1890s that he hoped would raise the status of the fruit. Two Burbank cultivars, `Van Deman' and `Pineapple', are important commercially in California today, but overall quince fruit production in the United States is so small that it is not even tracked by the USDA National Agricultural Statistics Service (McCabe 1996; USDA 2009b). While underappreciated here, these Burbank quinces have found their way to other parts of the world where they are among the handful of cultivars considered worthy of production (Campbell 2008). In 1908, Meech described 12 quince varieties important in the United States Burbank's `Pineapple' quince as seen in a photograph from the 1914 multi-volume publication Luther Burbank, His Methods and Discoveries and Their Practical Application. at the time, although some hat description hardly fits the quince known in America today, or rather the quince which is hardly known today. During Colonial times a quince tree was a rarity in the gardens of wealthy Americans, but was found in nearly every middle class homestead (Roach 1985). The fruit--always cooked--was an important source of pectin for food preservation, and a fragrant addition to jams, juices, pies, and candies. However, by the early twentieth century quince production declined as the value of apples and pears increased. Today's consumers prefer the immediate gratification provided by sweet, ready-to-eat fruits. After Charles Knox introduced powdered gelatin in the 1890s the use of quince pectin for making jams and jellies declined. U.P. Hedrick lamented in 1922 that T Quince 3 JOSEPH POSTMAN The attractive flowers and foliage of quince. like `Orange' (syn. = `Apple') were as often as not grown from seed rather than propagated as clones. Quince is easily grown from either hardwood or softwood cuttings, and is readily grafted onto another quince rootstock. Although it is an important dwarfing rootstock for pear, quince should not be grafted onto pear roots because this reverse graft is not reliable. Quince has a very extensive history in the Middle East, and may have even been the fruit of temptation in the story of the Garden of Eden. The ancient Biblical name for quince translates as \"Golden Apple\" and cultivation of Cydonia predates cultivation of Malus (apple) in the region once known as Mesopotamia, now Iraq. Juniper and Mabberly (2006) explain how this region is well adapted to cultivation of quince, pomegranate, and other fruits, but Mesopotamia was much too hot and dry for the cultivation of all but the most recently developed low-chilling-requirement apples. Quince was revered in ancient Greece, where a fruit was presented to brides on their wedding day as a symbol of fertility. It was mentioned as an important garden plant in Homer's Odyssey, and Pliny the Elder extolled its medicinal properties. Botany and Intergeneric Liaisons Cydonia oblonga is a monotypic genus belonging to family Rosaceae, subfamily Spiraeoideae, tribe Pyreae, and subtribe Pyrinae (USDA 2009a). It grows as a multi-stemmed shrub or small tree and has pubescent to tomentose buds, petioles, leaves, and fruit. Leaves are ovate to oblong, about 2 inches (5 centimeters) across and 4 inches (10 centimeters) long. The solitary white flowers are 1 to 2 inches (4 to 5 centimeters) across, have 5 petals, 20 or more stamens, 5 styles, an inferior ovary with many ovules, and are borne on current season growth. Bloom time overlaps with that of apples, usually beginning mid April in the central latitudes of the northern hemisphere. The fruit is a fragrant, many-seeded pome about 3 inches (8 centimeters) in diameter. Shape ranges from round to pear-like, flesh is yellow, and the Baileys refer to it as \"hard and rather unpalatable\" 4 Arnoldia 67\/1 Veitch in London sent scions of a quince-pear hybrid to Louis Trabut, the Algerian botanist. Trabut proposed the name Pyronia veitchii for this curious seedless-fruited hybrid (Trabut 1916). Pyronia is little known today, except by fruit tree pathologists who use the virussensitive clone as a graft-inoculated indicator to detect virus diseases in pome fruits. Another more recent hybrid generated in Japan between Cydonia and the Japanese pear, Pyrus pyrifolia, was probably the product of embryo rescue, a controlled tissue culture technique. In Italy and the Czech Republic, a purported hybrid between quince and apple (Cydomalus) has been touted as a possible rootstock for both apples and pears (Wertheim 2002). Illustration of `Orange' quince from U.P. Hedrick's 1922 Cyclopedia of Hardy Fruits. JOSEPH POSTMAN Center of Origin Cydonia is native to western Asia, and the center of origin is considered to be the Trans-Caucasus region including Armenia, Azerbaijan, Iran, southwestern Russia, and Turkmenistan (USDA 2009a). During ancient times, quince spread from its wild center of origin to the countries bordering the Himalaya Mountains to the east, and throughout Europe to the west. It has many uses and traditions associated with it throughout this broad range. Several recent USDA funded plant collecting expeditions to Armenia, Georgia, and Azerbaijan returned with quince seeds and cuttings from these countries. The availability of Cydonia germplasm in the United States increased significantly from 2002 to 2006 as a result of these collections (McGinnis 2007). Cultivation for Fruit and Rootstock Production Worldwide, there are about 106,000 acres (43,000 hectares) of quince in production with a total crop of 335,000 metric tons. Turkey is the largest producer with about 25% of world production. China, Iran, Argentina, and Morocco each produce less than 10%. The United States is a very minor player in quince fruit production with only about 250 acres (about 100 hectares) planted, mainly in California's San Joaquin Valley. Burbank's `Pineapple' is the most widely grown cultivar in that state and is said to be more flavorful than `Smyrna' (McCabe 1996). Quince fruit has a number of culinary uses. Dulce de membrillo, or quince paste, is popular in several European countries, particularly Spain. It is also much appreciated in parts of A Pyronia fruit--from a cross of Pyrus pyrifolia (Japanese pear) and Cydonia oblonga--growing in the USDA genebank orchard. (Bailey and Bailey 1976; Rehder 1986). Fruit size and leaf size of cultivated varieties can be many times larger than the wild type described above. All varieties are self-pollinating. Intergeneric crossing is fairly rare in plants, but has occurred naturally on occasion in the Rosaceae. While not as promiscuous as its cousins Sorbus and Mespilus, Cydonia has had a number of encounters with related genera that resulted in intergeneric offspring. In 1913 a Mr. Quince 5 JOSEPH POSTMAN Quince A will be about half the size of a tree grafted onto pear seedling rootstock. The tree will also be more precocious and fruit size will be larger. Quince C produces a tree slightly smaller and more precocious still. Provence Quince rootstock produces a pear tree slightly larger than Quince A or C. Some pear varieties are not graft compatible with quince and require a compatible interstem pear variety such as `Comice', `Old Home', or `Beurre Hardy' as a bridge. Dr. Vagharshak Hayrapetyan, head of the Scientific Center for Viticulture, Fruit Growing, and Winemaking in Yerevan, Armenia, poses with the winter quince variety `Chartar Gyugh' in September, 2006. Scions of this heirloom quince cultivar were recently brought to the United States. Landscape Use Few small trees rival the quince in becoming interestingly gnarled and twisted with age. Nonetheless, renowned Arnold Arboretum horticulturist Donald Wyman (Wyman 1965) STEPHEN AUSMUS, USDA ARS Latin America. This sweet, fragrant, jellylike confection is cut into slices and often served with a heady cheese. Quince is also served poached in either water or wine, and when so prepared develops a rich aroma and deep caramel-red color. In Armenia, quince is used in many savory as well as sweet dishes, and is often cooked with lamb (Ghazarian 2009). Quince fruit is also used by some home brewers to make very fine hard ciders. While quince is still grown for its fruit in some parts of the world, in England, France, and the United States it is primarily grown for use as a dwarfing pear rootstock. In the region around Angers, France, quince has been used as a pear rootstock since before 1500. The French were growing quince plants from cuttings and layering in stool beds by the early 1600s and France became an important source of rootstocks around the world. Quince rootstocks grown near Angers were known as `Angers Quince' and those propagated near Fontenay were known as `Fontenay Quince' (Roach 1985; Tukey 1964). Confusion arose about the identities of various quince rootstocks, and in the early 1900s researchers at East Malling in England collected rootstocks from a number of nurseries and designated clones with letters of the alphabet. Quince rootstock clones now available in the United States include Quince A and Quince C, which came from East MallingLong Ashton (EMLA); and Provence Quince (= Quince BA 29-C) from France. A pear tree grafted onto These bowls of quince show the diversity of shapes found in quince fruit. 6 Arnoldia 67\/1 JOSEPH POSTMAN This young quince tree, growing in the genebank orchard at USDA-ARS, Corvallis, Oregon, has been pruned to open up the crown and remove basal suckers. did not consider Cydonia worthy of his list of recommended landscape trees. He relegated it to his secondary list because of inferior flower interest, poor growth habit, and pest problems. However, Cydonia is an essential component of many historic gardens, and Frederick Law Olmsted included the common quince as a valuable plant in some of his landscapes (Deitz 1995). As a young tree, Cydonia may sucker profusely, and it takes some pruning effort during the first few years to establish an open-crowned specimen tree rather than a small thicket. Quince is such an interesting plant that it's worth the pruning effort, and germplasm recently imported from other parts of the world may provide some relief from pest and climate challenges that limited its use in the past. When grown in high pH soils, however, trees can become stunted and suffer iron chlorosis. In northern latitudes or colder climates the fruit of many cultivars does not fully ripen prior to the onset of winter, and in places where it rains during the ripening season, fruit cracking can be a big problem. Although most commercial quince production today is located in very warm areas, one of the largest quince orchards in 1895 was a 60 acre (24 hectare) planting in upstate New York near Waterport (Brown's Berry Patch 2007). Whether grown for fruit production or for use as a pear rootstock, quince is impacted by several disease problems. Fire blight caused by the bacterium Erwinia amylovora limits the cultivation of quince either for its fruit or as a pear rootstock, especially in regions with warm, humid summers. The genus Cydonia is one of the most susceptible to fire blight in Rosaceae, the plant family which includes many susceptible hosts (Postman 2008). Leaf and fruit spot caused by Fabraea maculata (anamorph = Entomosporium mespili) can result in tree defoliation and production of disfigured, unmarketable fruit if not controlled. Powdery mildew and rust diseases also impact quince production. Genetic improvements needed for expanding the use of quince as a dwarfing pear rootstock include increased resistance to fire blight for warm and humid summer climates, and increased winter cold-hardiness for northern climates. Adaptation to alkaline soils will allow quince production to expand to more diverse JOSEPH POSTMAN Potential for Genetic Improvement Quince is adapted to hot, dry climates and to acid soils. Under favorable conditions, ripe fruit can become quite fragrant, juicy, and flavorful. The Turkish cultivar `Harron' has the largest fruit size of the hundred or so quince clones growing at the USDA genebank, but the fruit may crack badly when exposed to rain just before it is ripe. Quince 7 JOSEPH POSTMAN A young boy in Georgia's northeast province of Kakheti displays quince fruit from a tree in the village of Shilda. Scions of the Shilda quince were collected by ARS genebank curators Joseph Postman and Ed Stover during a 2006 expedition to the Caucasus region. A tree is growing in quarantine at Beltsville, Maryland, and will be sent to the USDA-ARS genebank in Oregon upon release soil conditions both as a rootstock for pear or for production of quince fruit. Very slight progress in soil adaptation was achieved by selecting somoclonal variants of rootstock clone Quince A following multiple generations of in vitro culture on high pH media (Bunnag et al. 1996). Quince for fruit production will benefit from earlier ripening, and elimination of summer \"rat-tail\" blooms, which predispose a tree to attack by fire blight. Fruits that are picked too green will never ripen properly (McCabe 1996). Resistance to the fungal rusts and mildews will allow quince to be produced with fewer pesticide applications. Available Germplasm A quince germplasm collection was established in Izmir, Turkey, beginning in 1964 that includes many regionally developed fruit cultivars and landraces (Sykes 1972). In Karaj, Iran, a collection of more than 50 Cydonia accessions are maintained, including both cultivated and wild types (Amiri 2008). Smaller quince collections are growing in Italy, Greece, Spain, and other European countries (Bellini and Giordani 1999). There are also significant collections in Ukraine and southwest Russia. A large fruit tree collection in Kara Kala, Turkmenistan, was once a part of the Vavilov Institutes during Soviet times. Many fruit tree accessions, including quince, were rescued from that station in the late 1990s and brought to other genebanks for safekeeping. More than a dozen quince accessions from Kara Kala, representing both wild types and fruiting cultivars, are growing at the USDA genebank in Oregon. The Oregon facility is one of several ex situ genebanks housing temperate fruit and nut collections for the USDA National Plant Germplasm System (NPGS) (Postman et al. 2006). The NPGS Cydonia collection includes more than 100 clones with origins from 15 countries maintained as self-rooted trees in a field collection (Postman 2008). About half of this collection represents cultivars for fruit production, and the other half are pear rootstock selections, wild types, and seedlings. Observations made at the genebank have revealed a wide diversity of genotypes, some with resistance to Fabraea leaf and fruit spot, and a range of ripening seasons that may make it possible to produce quince fruit in short-season production areas. Quince selections made in Bulgaria following a fire blight epidemic in that country have shown good field resistance to the disease, and some of this Bulgarian germplasm was recently introduced into the United States by the NPGS genebank. For nearly a century, the quince has been almost ignored for fruit production in North America, while many improvements have been made in the Middle East and central Asia. Germplasm is now available in the United States for expanding the use of Cydonia both as a rootstock for pear and as a fruit producing tree in its own right. As Luther Burbank concluded a hundred years ago, \"The quince of today is, indeed, a half wild product that has waited long for its opportunity. It remains for the fruit growers of tomorrow ... to see that the possibilities of this unique fruit are realized\" (Wickson et al. 1914). 8 Arnoldia 67\/1 his Chinese relative of Cydonia presently belongs to the genus Pseudocydonia, but has previously been assigned to both Chaenomeles (Chaenomeles sinensis) and Cydonia (Cydonia sinensis). Chinese quince has attractive single pink flowers that appear earlier than those of Cydonia but not as early as most Chaenomeles. The fruit is a large, oval, aromatic yellow pome that ripens in the fall. The shiny, leathery leaves develop nice red-orange fall color. But its most interesting characteristic is the exfoliating bark that reveals brown, green, orange, and gray patches. Chinese quince's attractive bark rivals that of many stewartias. The trunk often becomes fluted with age, adding even more textural appeal. Luther Burbank devoted some attention to the Asian quinces and was probably responsible for a large-fruited clone of Pseudocydonia. Michael Dirr (1997) notes that Chinese quince is reliably hardy in USDA Zones 6 to 7 (average annual minimum temperatures -10 to 10F [-23 to -12C]), and possibly hardy in Zone 5 (-20 to -10F [-29 to -23C]). Fire blight is said to seriously impact its cultivation. However, the presence of very nice specimens of Chinese quince at the National Arboretum in Washington, D.C., and in gardens in the Carolinas--locations where Cydonia is readily killed by fire blight--indicate that it can be grown even in regions where the disease is present. Chinese quince's pink flowers, attractive patchwork bark, and fluted trunk are highly ornamental. COURTESY OF KENPEI MICHAEL DOSMANN T The Chinese Quince: Pseudocydonia sinensis Quince 9 References Amiri, M.E. 2008. The status of genetic resources of deciduous, tropical, and subtropical fruit species in Iran. Acta Horticulturae 769:159167. Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third. Campbell, J. 2001. Quince Growing. New South Wales AgFact H4.1.3. Bellini, E. and E. Giordani 1999. Online European Minor Fruit Tree Species Database EMFTS Database. http:\/\/www.unifi.it\/project\/ueresgen29\/ netdbase\/db1.htm (7 March, 2009). Brown's Berry Patch. 2007. http:\/\/www.brownsberrypatch. com\/history_farm.html (2 April, 2009). Bunnag, S., R. Dolcet-Sanjuan, D.W.S. Mok, and M.C. Mok. 1996. Responses of two somaclonal variants of quince to iron deficiency in the greenhouse and field. Journal of the American Society of Horticultural Science 121:10541058. Deitz, P. 1995. Fairsted: at home with Frederick Law Olmsted. Magazine Antiques, August 1995. Dirr, M.A. 1997. Dirr's Hardy Trees and Shrubs, An Illustrated Encyclopedia. Timber Press, Portland, Oregon. Ghazarian, B. 2009. Simply Quince. Mayreni Publishing, Monterey, CA. 216 pp. Hatch, P.J. 1998. The fruits and fruit trees of Monticello. University Press of Virginia. pp. 127128. Hedrick, U.P. 1922. Cyclopedia of Hardy Fruits. Juniper, B.E. and D.J. Mabberly. 2006. The story of the apple. Timber Press, Portland, OR. 219 pp. McCabe, C. 1996. Enjoying the forbidden fruit. Saveur 14:105110. McGinnis, L. 2007. Quest for Quince: Expanding the NCGR Collection. Agricultural Research, January 2007:2021. Meech, W.W. 1908. Quince Culture; an illustrated handbook for the propagation and cultivation of the quince, with descriptions of its varieties, insect enemies, diseases and their remedies. Orange Judd Co., New York. 180 pp. Postman, J. 2008. The USDA Quince and Pear Genebank in Oregon, a World Source of Fire Blight Resistance. Acta Horticulturae 793:357362. Postman, J., K. Hummer, E. Stover, R. Krueger, P. Forsline, L.J. Grauke, F. Zee, T. Ayala-Silva, B. Irish. 2006. Fruit and Nut Genebanks in the US National Plant Ger mplasm System. HortScience 41(5):11881194. Rehder, A. 1986. Manual of Cultivated Trees and Shrubs Hardy in North America, 2nd edition. Dioscorides Press, Portland, OR. Rieger, M. 2006. Mark's Fruit Crops. http:\/\/www.uga. edu\/fruit (4 February, 2009). USDA. 2009a. ARS, National Genetic Resources Program. Germplasm Resources Information Network (GRIN) [Online Database]. URL: http:\/\/www. ars-grin.gov\/cgi-bin\/npgs\/html\/taxon.pl?12779 (20 January 2009) Roach, F.A. 1985. Quinces. In: Cultivated Fruits of Britain: Their Origin and History. Blackwell, London pp. 220225. Sykes, J.T. 1972. A description of some quince cultivars from western Turkey. Economic Botany 26:2131. Trabut, L. 1916. Pyronia: A hybrid between the pear and quince. Journal of Heredity 7:416419. Tukey, H.B. 1964. Dwarfing rootstocks for the pear. Ch. 11 In: Dwarfed Fruit Trees, The MacMillan Co., New York. pp. 182199. USDA. 2009a. Germplasm Resources Information Network - (GRIN) Online Database. National Germplasm Resources Laboratory, Beltsville, Maryland. http:\/\/www.ars-grin.gov\/cgi-bin\/ npgs\/html\/taxon.pl?12779 (05 Febr uar y 2009) USDA. 2009b. National Agricultural Statistics Service, U.S. fruit production data. http:\/\/www.nass. usda.gov\/QuickStats\/indexbysubject.jsp (4 February, 2009) Wertheim, S.J. 2002. Rootstocks for European pear: a Review. Acta Horticulturae 596:299309. Whitson, J., R. John, and H.S. Williams (eds.) 1914. The Transformation of the Quince. Chapter 7, Volume 4 In: Luther Burbank, His Methods and Discoveries and Their Practical Application. Luther Burbank Press, New York and London pp. 211240. Wyman, D. 1965. Trees for American Gardens. Macmillan Publishing Co., New York. Joseph Postman is a plant pathologist and curator of living pear, quince, and hazelnut collections at the USDA Agricultural Research Service, National Clonal Germplasm Repository in Corvallis, Oregon. Information about quince genetic resources in the USDA National Plant Germplasm System, as well as a field day on October 10, 2009 featuring the Corvallis genebank's quince orchard, is available at http:\/\/www. ars.usda.gov\/pwa\/corvallis\/ncgr. A one day symposium on underutilized pome fruits will be held in August, 2010 during the 28th International Horticulture Conference in Lisboa, Portugal. For more information visit the `Symposia' link at http:\/\/www.ihc2010.org\/ "},{"has_event_date":0,"type":"arnoldia","title":"Bird's-eye Views: Aerial Photographs of the Arnold Arboretum","article_sequence":2,"start_page":10,"end_page":19,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25460","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14e856d.jpg","volume":67,"issue_number":1,"year":2009,"series":null,"season":null,"authors":"Connor, Sheila","article_content":"Bird's-eye Views: Aerial Photographs of the Arnold Arboretum Sheila Connor ur desire to fly must have been driven, in part, by wanting to have a bird's-eye view of the land. Today, we can launch ourselves skyward simply by clicking on Google Earth, where a virtual world created by combining aerial photography, satellite imagery, and GIS (geographic information systems) unfolds on our computer screen. Attainment of that instant bird's-eye view was many years in the making, though. The first aerial photographs were taken from a hot air balloon in 1858 by the French portraitist \"Nadar\" (GaspardFelix Tournachon), who did so while tethered 240 feet (73 meters) above the village of Petit-Bicetre near Paris. Two years later--also in a tethered hot air ballon-- James Wallace Black ascended 1,200 feet James W. Black's 1860 image of Boston, the earliest aerial photo still (366 meters) over the densely developed in existence. port city of Boston, Massachusetts. His the shutter placed inside the lens, his highimage, \"Boston, as the eagle and wild goose see speed camera was capable of producing images it,\" is the earliest known aerial photograph still with little or no distortion, which made accuin existence. Kites, rockets, and carrier pigeons rate mapping possible. Although the army did (outfitted with tiny breast-mounted cameras) not take delivery of his cameras until after the were the next airborne means used. war, Fairchild continued to improve upon his Just a few years after the Wright brothers design and, in 1920, started the Fairchild Aerial famous first flight, images were shot from an Camera Corporation. airplane piloted by Wilbur Wright, the first He also began designing aircraft to suit his taken from an airplane. The military, both photographic needs and founded his second here and abroad, quickly grasped the value of company, Fairchild Aerial Surveys, Inc. The these unexpectedly revealing views and estabcompany is well known for the remarkable lished aerial reconnaissance units. Following aerials it produced of every major city in the World War I, newly created commercial comUnited States between 1920 and 1960, and panies expanded upon the progress made in the Arboretum was one of its earliest clients. aerial techniques. Using one of the company's specially designed New Equipment, New Techniques cameras, a pilot flew over the Arboretum in Sherman Mills Fairchild started several of 1927 in a Fairchild FC-1 and took \"the first these new peacetime ventures. Fairchild had airplane view to show all of America's greatoriginally secured a contract with the army to est hardy garden,\" as reported in the Boston develop a camera for aerial photography. With Herald newspaper. This \"bird's-eye view\" was O COUrTESy OF THE BOSTON PUBlIC lIBrAry Aerial Photographs 11 This 1927 Fairchild aerial photograph of the Arboretum, looking toward Boston, shows Peters Hill in the foreground and the familiar curlicue of roadway atop Bussey Hill. JAy CONNOr This 2005 image was made with the same perspective as the 1927 photo. Peters Hill is in the foreground, but mature trees now obscure the top of Bussey Hill. Downtown Boston is seen in the distance. All PHOTOS FrOM THE ArCHIvES OF THE ArNOld ArBOrETUM UNlESS OTHErWISE INdICATEd 12 Arnoldia 67\/1 A large paved circle for bus turnarounds is seen atop Peters Hill in this 1967 photograph. Prior to 1964 there was no paved roadway to the top of the hill. In the late 1990s the paving was removed as part of a landscape restoration project that returned the hilltop to a design consistent with Frederick Law Olmsted's naturalistic style. JAy CONNOr reproduced in the the newspaper's autogravure section on November 20, 1927. Since that initial flight, photographers have used planes, helicopters, a dirigible, and, most recently, a drone as means to attain views of the Arboretum. The resulting collection of negatives, microfiche, prints (both black and white and color), and digital images provides a unique perspective and an amazing record of how change occurs in the Arboretum's seemingly permanent landscape. Entire plant collections disappear only to reappear years later completely redesigned and reconfigured. Others simply disappear. A few migrate, acquire a new name, then eventually vanish. roads appear, are paved, then unpaved, and fade away. Sidewalks and paths (whether planned or established by desire) do the same, and while our aerial archaeology has not revealed any crop circles, one can easily see the remains of the characteristic circles that signify abandoned planting holes, sites where specimens once grew. Additional unplanned footpaths created over the years are visible in this 2007 image of Peters Hill. In place of the pavement at the summit there are now a scattering of granite blocks used for informal seating. The granite blocks, recycled from a demolished Olmsted-era bridge that once stood near the Forest Hills Station, were originally placed in a circle on Peters Hill in the 1980s to deter a then popular youth activity--setting stolen cars on fire and pushing them down the hill. Making Maps The first vertically shot aerial survey of the living collections took place in 1936. (In vertical aerial photography the camera is in a level position and pointing directly downward, the best format for precise mapping.) This survey consisted of a series of four images taken by Bradford Washburn, then a 26-year-old instructor at Harvard's Institute for Geographical Exploration. His long Aerial Photographs 13 When seen from above in this 1955 Bradford Washburn aerial (top), the broad, grassy plain just below the summit of Bussey Hill sports shadows of planting holes from the Prunus collection that once occupied the site, seen in the May 1929 photo (bottom) taken by the renowned New England landscape photographer Herbert Wendell Gleason. affiliation with the Arboretum, coupled with his expertise in aerial photography and cartography, greatly influenced the number of aerial photographs taken of our landscape. Mr. Washburn often acted as a project manager, directing and organizing both vertical and oblique (camera is angled) shots made of the arboretum. Under his direction an image of the 14 Arnoldia 67\/1 1 2 3 7 6 4 5 One of Washburn's 1936 vertically shot aerials of the Arboretum. Marked on the map are: 1. Present site of the Dana Greenhouse, constructed in 1962, and the Leventritt Pavilion and Shrub and Vine Garden (an aerial view of this garden is on the front cover). 2. The site of the original Shrub and Vine Collection, now occupied by the Bradley Rosaceous Collection. 3. Site of the Bussey Institution, the location of the Arboretum's greenhouses prior to 1962, and now the site of the Massachusetts State Laboratory. 4. Bussey Brook Meadow, also known as the South Street Tract and Stony Brook Marsh, prior to the pond being filled in and the creation of the Blackwell Footpath 5. Peters Hill had only the outer ring road at the time. 6. Weld Hill, once known as the Weld Walter Street Tract, prior to the construction of the Hebrew SeniorLife Center on the site of the former Joyce Kilmer Park 7. Highly visible remnant of Centre Street left from the Centre Street realignment and widening in 1931. Today, a grassy swath still indicates the route of the old roadbed. Aerial Photographs 15 entire Arboretum was taken in 1952. Then in 1955, his first year as chairman of the Arboretum's visiting Committee, he raised the sum of $310.00 from the committee for a flyover by Eastern Aerial Surveys, Inc., with the recommendation that a second survey take place the following spring. Twelve images resulted from the October 6 survey. Unfortunately there is no record of a spring session. Northeast Aerial Photos produced the first series of color images of the Arboretum in 1967. A year later, color images of the Hunnewell Building and the newly built garage facility were taken, and in 1974 a survey of the entire Arboretum produced a suite of seventeen images. Bradford Washburn's longheld goal of creating a mapping system of the Arboretum's living collection based on aerial photography finally came to fruition when dr. Peter Shaw Ashton, then director of the Arboretum, approached him in 1978 to orchestrate the coordination of a photogrammetric survey of the Arboretum by Swissair Photos + Surveys, ltd. (now named Swissphoto AG). \"On a cloudless day in April, 1979, the survey crew took a series of aerial photographs, which were then transformed into orthographically corrected images displaying an exceptionally accurate picture of the Arnold Arboretum at a scale of 100 feet to the inch. A groundsurvey team was hired to complete the contours in certain areas of the Arboretum that are covered by an evergreen canopy. Swissair provided the Arboretum with a base map of the grounds that illustrates true north, contour lines at intervals of ten feet, physical features (roads, paths, walls, and buildings), and reference points.\" From the article \"Cartographic records of the living Collections\" Ethan W. Johnson, Arnoldia. 49 (1) 1989. The Hunnewell Building and then newly built garage behind it are shown in this 1968 photograph. JAy CONNOr A similar view as seen in May, 2005. 16 Arnoldia 67\/1 B rAdFOrd WASHBUrN was an extraordinary man. Born in Boston in 1910, as a teenager he developed a love for mountain climbing, summiting peaks around the world in the days well before high tech climbing gear was available. As an undergraduate at Harvard he honed his passions--climbing, photography, and scienctific exploration-- and in 1934 pursued graduates studies in cartography, surveying, and aerial photography at Harvard's Institute for Geographical Exploration. At 29 he became the director of the New England Museum of Natural History, now the Boston Museum of Sci ence, a position he held for 40 years. As a pioneer in aerial photography, Washb u r n 's s t u n n i n g mountain images made him one of the most important landscape photographers of the twentieth century. recently, one of Washburn's cameras (a 1929 Zeiss 4 x 5) was taken on Bradford Washburn and the Fairchild 71 Monoplane, Valdez, Alaska, a the space shuttle's 1937 gelatin silver print photograph by Bob Reeve. Hubble telescope repair mission by astronaut and mountain climber John Grunsfeld. It seems fitting that Washburn's camera was used to make the ultimate in aerial photos--images from space. PHOTOGrAPH 2009 MUSEUM OF FINE ArTS, BOSTON. GIFT OF BrAdFOrd ANd BArBArA WASHBUrN. 1999.446 Aerial Photographs 17 Susan Kelley, then curatorial associate in mapping, and I met with Mr. Washburn in 2000 to learn more about his early Arboretum work. He believed that his photographs of the collections would eventually provide a valuable record. Upon seeing our current maps of the living collections, which were based on the 1979 photogrammetric survey and formatted in AutoCAd, which interacts with the computerized plant records database, BG-Base, Mr. Washburn pronounced them \"gorgeous!\" More Bird's-eye Views Sasaki Associates ltd. produced aerials in 1990 and 1991 as part of the Arboretum's Master Plan process, and in 2002 the Arboretum participated for the first time in a survey of the Harvard campus, which was coordinated by Harvard's Planning and real Estate department. The living collections were again included in the Harvard survey in 2006. recently, when aerial imagery has been needed, photographs have also been acquired from surveys done by the United States Geological Surveys (USGS). Our most recent full scale vertical aerial view of the entire Arboretum was taken in spring 2008, as part of the USGS Boston 133 Cities Urban Area mapping program. Interspersed between these major surveys were other more site specific or overview flights. In 1950, Arboretum horticulturist donald Wyman took a series of photographs at a Image of the Arnold Arboretum in 2008 from the United States Geological Surveys. 18 Arnoldia 67\/1 Weld Hill in 1955, with Kilmer Park adjacent (top of photo). Weld Hill in 2006, part of the Harvard survey. height of 3,000 feet from a plane flown by his 17-year-old son.Wyman's photographs were taken from a vantage point reminiscent of the 1927 Fairchild survey images. Eight years later, Heman Howard, in charge of the mapping and labeling department, also duplicated this view with a series of oblique shots from both 1,400 and 2,300 feet. The Massachusetts department of Public Works photographed the lilac collection and replicated the bird's-eye view in 1969 and, in 1995, Sergio Marino of GPI Models took a series of images from a helicopter to facilitate his construction of an 8 feet by 16 feet scale model of the Arboretum. The model became Aerial Photographs 19 An oblique view of Weld Hill, taken from a drone on May 20, 2009, shows the Arboretum's new research facility under construction. the centerpiece for the exhibit Science in the Pleasure Ground in the Arboretum's Hunnewell visitor Center, where it continues to be a popular feature. My brother, Jay Connor, has taken almost 200 oblique images of the collections. He began photographing the Arboretum in 2004, usually from a helicopter, but once from the iconic Hood Blimp, officially an American A-60+ lightship. This familiar cigarshaped balloon is capable of hovering motionless for hours at a time. As Boston red Sox fans can attest, it is truly an airship designed for aerial observations. The Arboretum's most recent aerial photography project involves the new research facility on Weld Hill. Over the past two years, dave Fuller of Fullerview Photography has sent up a drone to capture images of the construction of the building. Over the coming months, arboretum staff will be adding aerial imagery to our GIS (geographic information system) using ESrI software. This will provide a new generation of bird's-eye views of the arboretum's landscape and its change over time. Incorporating these images into our GIS system will assist in reconciling diverse georeferenced features and provide unprecedented detail about our living collections for researchers and visitors. Sheila Connor is the Horticultural research Archivist at the Arnold Arboretum. FUllErvIEW PHOTOGrAPHy "},{"has_event_date":0,"type":"arnoldia","title":"Seeking Cold-Hardy Camellias","article_sequence":3,"start_page":20,"end_page":30,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25465","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15ea76d.jpg","volume":67,"issue_number":1,"year":2009,"series":null,"season":null,"authors":"Aiello, Anthony S.","article_content":"Seeking Cold-Hardy Camellias Anthony S. Aiello or those of us in more northern climates, trips to southern or West Coast gardens in early spring often result in admiration (and a little envy) for the range and beauty of camellias (Camellia spp.) that can be grown in Zones 7 or warmer. As with many plants, we always want those that are either too tender or too boreal for our zone; those plants well suited for a particular climate are all too quickly considered prosaic and it is the struggling arcane plants that most of us cherish as gardeners. It was the tantalizing possibility of finding more cold-hardy camellias that 25 years ago led to a plant hunting expedition and the resulting multi-year evaluations of a group of Camellia japonica. Domestic and international plant exploration, and subsequent evaluation of plant acquisitions have been important missions of the Morris Arboretum in recent decades. Since the late 1970s, staff of the Morris Arboretum have participated in 20 plant collecting trips, including trips to South Korea, China, the Caucasus Mountains, and regions within the United States. On these expeditions, seed is collected and returned to the Morris Arboretum for propagation. (Occasionally live plants are collected, but because of difficulties with transportation and import regulation, seeds are the primary form collected.) One of the main goals of our plant exploration and evaluation program is broadening the genetic pool of known species to extend cold hardiness and increase vigor. Between 1979 and 1991, Morris Arboretum staff participated in five collecting expeditions to South Korea. These trips were planned to sequentially cover different geographic regions of South Korea. The 1984 Expedition to Korea's northwestern coast and islands (Korea Northwest Expedition KNW) visited areas along the northwestern coast and inland to the Kwangnung Arboretum (now Korea National Arboretum) of South Korea (Meyer 1985). It is from this 1984 expedition that the Morris holds a number Map of areas visited on the 1984 Korea Northwest collecting expedition. of accessions of Camellia japonica collected on Taechong and Sochong Islands, off the west coast of South Korea. The island collections represent some of the most northern collections ever made of common camellia. As an extension of the Asian land mass, Korea is exposed to a continental climate that includes strong, cold, and persistent winter winds. Even along the coast, the Korean climate is much harsher than that in Japan. As a result, despite their location in the Yellow Sea, these islands are exposed to more extreme temperatures than one would expect from their maritime location. The Trip to the Islands The idea of visiting and collecting from these island populations of Camellia japonica was instigated by Barry Yinger (Asiatica Nursery, MOrrIS ArBOrETUM OF ThE UNIvErSITY OF PENNSYlvANIA F Camellias 21 PAUl W. MEYEr The Taean Peninsula on the northwest coast of South Korea. This area and islands off of the northwest coast were the focus of the 1984 Korea Northwest collecting expedition. lewisberry, Pennsylvania), who had read of this northern cold-hardy population in the early 1980s (Yinger 1989a; Yinger 1989b). Through great persistence, Yinger first encountered these plants on Taechong and Sochong Islands in the winter of 1981. Yinger relates how his concern that the camellias were destroyed during the almost total deforestation of Korea during World War II turned to delight once he reached the islands. Yinger wrote about his first encounter with these camellias on Sochong Island: \"... off we went, up the hillsides overlooking the Yellow Sea, buffeted by the cold wind from the northeast. The hillside was bleak and brown with few trees of any kind. The only greenery was an occasional grove of pines, the lower limbs of which had been chopped off for firewood. Up a little further and there--at last--a grove of camellias glittering green against the brown dried grasses, catching the winter sunshine and throwing it back to us.\" (Yinger 1989a) By counting the growth rings of stumps of camellias that were cut for firewood, Yinger estimated the age of these trees, some of which were 15 to 18 feet (4.6 to 5.5 meters) tall, as close to 150 years old. These astonishing trees had silently witnessed the political vagaries that had affected the Korean peninsula and its people over that long period. In October 1984, Yinger, then at the U.S. National Arboretum, returned to Taechong, Sochong, and Paekryong Islands with Sylvester March, Paul Meyer, and Peter Bristol (of the National, Morris and holden Arboreta, respectively) along with their Korean colleagues Chang Yong June and Chang Yong hun. Although these islands are controlled by South Korea they are located just south of the 38th parallel and north of the mainland border between North and South Korea. The islands are within view of North Korea, so they are of military and political significance; the explorers were required to 22 Arnoldia 67\/1 PAUl W. MEYEr Mature plants of Camellia japonica growing on a steep hillside on Sochong Island. Peter Bristol and Sylvester March are pictured at lower left. have a naval escort to reach and travel on the islands and were forbidden from photographing the boat on which they travelled. As Meyer (1985) wrote: \"... it must have been a peculiar sight as the Korean navy boat pulled out of Inchon harbor. Among the Korean sailors were four American plant explorers eager to collect on a group of islands in the Yellow Sea. Piled high on the deck were herbarium presses, seed bags, and general expedition supplies. The pole pruners leaning against the gun turrets created a strange juxtaposition. If the North Koreans observed this they must have wondered what this unusual mission was all about.\" Although the collecting supplies were exposed to the sea air, the Americans were sequestered below decks in crowded cabins for the duration of the long trip. Once on the islands, the collectors were escorted by the sailor companions, who eventually chipped in and helped with seed collecting and cleaning (trip details from Yinger 1989a and 1898b; Meyer 1985; and Meyer, personal communication). The Americans travelled among the three islands for approximately one week, making a large number of collections from a great diversity of plants. Among these were nine seed collections of Camellia japonica including some Camellias 23 PAUl W. MEYEr Collecting seeds from mature, open-grown Camellia japonica plants on Sochong Island. An unidentified Korean sailor is standing beneath the trees at left. that were growing in pastures and others that had been transplanted into local farmers' gardens. Six of these came from Taechong Island and the other three from Sochong Island. The islands' inhabitants recognized the beauty of these plants and often transplanted them into their small home gardens. Meyer (1985) found a grove on Sochong Island to be the most impressive; here, the camellias grew into large trees that grew luxuriantly on a site exposed to sea winds and salt spray. The areas where they grew were heavily cut and grazed by goats. Only tall plants with their lower foliage eaten remained, and the grazing prevented any natural regeneration of seedlings. (here at the Morris we have a similar problem, except it is the white-tailed deer that browse on our low hanging camellia foliage.) The human and livestock pressure on the islands was significant and the field notes describe collecting from resprouting plants Fruit of Camellia japonica collected on the 1984 Korea Northwest expedition. The camellia fruit is a woody capsule containing several seeds. in locations that were either cut-over forests, heavily grazed, or along roadsides. As unromantic as these types of plants and locations may sounds, they can make for excellent field PAUl W. MEYEr 24 Arnoldia 67\/1 throughout the year. Another standout from this group is Lindera obtusiloba; anyone who knows the sublime golden yellow fall color of Japanese spicebush agrees that it is one of the most outstanding shrubs for autumn foliage. ArNOlD ArBOrETUM Wanted: A Hardy Camellia What was the impetus that led to such effort to reach a far-flung corner of the world? As mentioned previously, camellias are exquisite garden flowers, but the vast majority of camellia cultivars are not hardy in regions colder than USDA hardiness Zone 7. From the late 1970s into the early 1980s a series of extremely cold winters devastated camellia collections at the U.S. National Arboretum and elsewhere (Ackerman 2007; Ackerman and Egolf 1992). At the National Arboretum alone, the harsh winters reduced the collection of 956 The handsome foliage of Japanese spicebush (Lindera obtusiloba) in 30- to 40-year-old plants to less than a golden fall color. dozen struggling survivors (Ackerman collecting. Compared to a mature forest, with 2000; Ackerman and Egolf 1992). These severe little sunlight reaching the understory and winters--and the damage to large numbers of fruits far out of reach, roadsides or regrown cultivars--inspired Dr. William Ackerman, a areas provide plants with sufficient sunlight to plant breeder and camellia aficionado at the produce fruit while lending easy access to the National Arboretum, and Dr. Clifford Parks, a plant collector. professor from the University of North Carolina In addition to the camellias, numerous other in Chapel hill, to undertake breeding programs plants were collected on the islands, and many to develop camellias cold-hardy in Zones 6 and of these have grown exceptionally well for us. 7. In light of the severe winters at the time of Most notable among these collections are Calthe Korean expeditions, there was considerable licarpa japonica, Lindera obtusiloba, Sorbus excitement about the potential for cold-hardy alnifolia, Styrax japonica, Pinus thunbergii, provenances of Camellia japonica coming from and Viburnum bitchuense. Meyer (1985) was South Korea (Yinger 1989a). It was hoped that particularly impressed by seaside populations these northern collections of Camellia japonica of Styrax japonica, which were noteworthy would expand the hardiness of common camelbecause of leathery and glossy leaves that were lia, generally considered to be reliably hardy in unaffected by salt spray or summer sun. Plants Zone 7 (Flint 1997) but historically not reliably grown from this seed collection grace our parkcold hardy in the Philadelphia area (Zone 6b). ing lot where their May flowers provide a fraThe nine accessions of Camellia japonica grant welcome to our visitors. Over the years were collected on the Korean islands in October we have lost many compound-leaved Sorbus 1984, and some of these seeds were sown at the species, but perhaps the best mountain ash for Morris Arboretum beginning in November of our area is Sorbus alnifolia. With its simple that year. Eight of the nine accessions germileaves, abundant white flowers, striking coralnated successfully, with varying numbers of red fruits, and russet fall color, the Korean seedlings among accessions. Given the northern mountain ash is one of my favorite plants locations of the parent populations, we began Camellias 25 The Tryouts Begin In 1986 plants were designated for one of two parallel evaluation studies: either a replicated field trial, or garden settings throughout the Arboretum. Of the eight successfully germinated accessions, six were eventually planted in the Arboretum's trials or throughout the Arboretum. In April 1987, 730 seedlings were planted in a replicated field trial at the Arboretum's Bloomfield Farm research area and were evaluated for cold hardiness. From 1989 to 1993 all of these Field trials of Camellia japonica at the Morris Arboretum's Bloomfield plants were evaluated for general foli- Farm, February and April, 1994. age quality, vigor, and hardiness (survival) on a scale of 1 to 5 (with 1 being dead plants were moved to our greenhouses. Then, and 5 being excellent). As would be expected between the fall of 1995 and spring of 1999, with seedling grown plants, there was great 25 of these highest rated plants from the origivariation in the survival and quality of plants nal 730 in the Bloomfield trial were planted in this study (Aiello et al. 2008). into the Arboretum's public garden for further By June 1990, 589 plants survived, and 283 assessment (Aiello et al. 2008). were deemed acceptable because they had a ratIn a parallel study, between 1987 and 1991 ing of 3, with only slightly damaged foliage. an additional 33 of the originally germinated Three years later, in August 1993, the cutoff seedlings that were not part of the formal field for retaining plants was elevated to a 4 ranking, trial were planted in protected garden settings that is, plants that showed only occasional foliar throughout the Arboretum. These plants did damage. At this level of scrutiny only 40 of 170 not receive the formal ratings applied to their remaining plants made the grade. The winters siblings in the research plots. Nevertheless, the of 199394 and 199495 resulted in further winters took their toll and by October 1999, 22 loss of plants, and by April 1995 the remaining of these plants remained in the garden. rICK J. lEWANDOWSKI a long-term field and garden trial of several accessions. Since the late 1980s plants grown from these collections have been evaluated for cold hardiness and several ornamental characteristics such as general vigor, leaf quality and retention, flower abundance and color, and plant habit. The camellias in this study all exhibit attractive evergreen foliage and single red flowers, which is typical of the straight species. These plants are large shrubs, reaching up to 12 feet (3.6 meters) tall or higher in 25 years. Although their single red flowers are not like the very showy forms grown farther south, their greatest value is in their hardiness and potential for breeding. rICK J. lEWANDOWSKI 26 Arnoldia 67\/1 Bringing it All Together In October 1999, shortly after I arrived at the Morris Arboretum, a total of 50 camellias were alive in garden settings throughout the Arboretum. Faced with what was already a 15-year old trial, I wanted to bring some resolution to this evaluation effort and to determine which of the remaining plants truly stood out among the others. The 50 plants included the 25 plants from the field trials, 22 remaining plants from those originally planted in garden settings, and three additional plants which had been cutting-grown in our greenhouse from original seedlings. These 50 plants were growing in protected areas throughout the Arboretum, where Camellia japonica plants growing at Gates Hall at the Morris Arboretum. the camellias could grow under the year's ratings for these plants were combined. canopy of conifers or against buildings, where These 43 plants were grouped into three catthey were shielded from strong winter winds egories according to overall performance and and afternoon sun. For example, one group was appearance after 5 years of evaluation. These massed to the north of a very large Chamaecycategories were somewhat subjective but paris pisifera that screens our parking lot from allowed us to consolidate several seasons of Meadowbrook Avenue, a quiet residential street information into a shorthand that would clarify that borders our property. Another group was the better performing plants. planted along the northeast face of Gates hall, Of the 43 plants, the top 15 (\"A\" rating) the Arboretum's administration building. exhibited consistent, positive performance in Starting in the fall of 1999 and continuthree key areas of the evaluation criteria. In paring through the spring of 2004, the 50 plants ticular, these plants flowered every year, mainthroughout the Arboretum were visually evaltained a desirable habit, and retained attractive uated. In the spring and fall of each year the glossy green foliage throughout the seasons. plants were rated for a variety of ornamental The foliage quality is especially important in traits including general vigor, hardiness, leaf March, when the effects of winter start to show retention, and foliar and floral characterison poorer performing plants. Because Cameltics. Plants with foliage that was deep green, lia japonica flowers on old wood before new glossy, disease-free, and with no winter injury growth emerges, we were especially interested received the highest ratings. Although there in those plants that retained high quality foliage was not a great deal of variation in floral traits, as the flowers emerged from March into April. plants with greater numbers of flower, flowers The middle 16 plants (\"B\" rating) generally perthat were more open, and flowers with richer formed well in one or two areas of the evalubright scarlet color were considered the most ation, but their performance was either not desirable. There was also significant variaconsistent, or was poor in the other categories. tion in plant habit and we gave higher ratings For instance, \"B\" plants may have had good to denser and more regularly shaped plants foliage quality but their flowering was poor or (Aiello et al. 2008). inconsistent, or they might have had beautiful After these visual evaluations were completed flowers but scraggly open habits that detracted in late 2004, 43 plants remained alive and each ANThONY S. AIEllO Camellias 27 from the overall quality of the plant. The lowest rated 12 plants (\"C\" rating) generally performed poorly in several categories. In some instances, they may have exhibited one positive characteristic, but this was overridden by the overall appearance of plant. The Current Situation and Next Steps After more than 20 years of evaluation, the numbers of Korean Camellia japonica at the Arboretum has gone from approximately 750 plants to just over 40 individuals. The remaining plants represent six of the original nine collections from Korea (KNW 312, 342, 344, 348, 350, and 352) and are a valuable genetic resource for introduction and breeding. Although their ornamental value may not compare to cultivars hardy in the southern and western United States, our plants exhibit attractive single red flowers and glossy evergreen foliage. They rep- resent a significant advance in the hardiness of common camellia, with suitability for Philadelphia and the mid-Atlantic region, and possibly the lower Ohio valley and coastal New England. These cold-hardy selections will appeal to Zone 6 gardeners who have coveted these plants after visiting the \"Camellia Belt\" found in southeastern and West Coast states. Along with evaluating the remaining plants in our collection, over the past several years we have been propagating and distributing cutting-grown individuals from our highest rated plants. Camellias have been provided to other public gardens throughout the northeastern United States, including Chanticleer, and the Scott, Tyler, Willowwood, Polly hill, and Arnold Arboreta. Our hope is that distributing this material will help conserve the germplasm and provide evaluation over a broader range of climates. ANThONY S. AIEllO Camellias with glossy green foliage that remained attractive through the winter received higher ratings in the evaluation. 28 Arnoldia 67\/1 ANThONY S. AIEllO Single, red flowers were standard for the Korean seedlings, though some plants had more vibrant color or greater numbers of flowers. Currently we are planning to name and introduce several individual plants from our Camellia japonica trials (see sidebar). Two of these plants are those that show the highest ratings for combination of plant habit, foliar quality, and flower density. One plant shows a striking upright habit and a fourth is consistently precocious, regularly blooming in late autumn compared to the normal early spring blooming time of the species. Presently there are three commercially available introductions from the 1984 Korean Camellia japonica collections. These are: `Korean Fire' (KNW 352) a 2003 Pennsylvania horticultural Society Gold Medal winner that was introduced by Barry Yinger through hines Nursery (Bensen 2000); and `longwood valentine' and `longwood Centennial' (KNW 350) introduced by longwood Gardens (Tomasz Aniko, personal communication). Going forward, our goal is to distribute our selections and compare them to other known cold-hardy forms of Camellia japonica. We are also working with plant breeders to share material in the hope that the hardiness inherent in our plants can be utilized to develop coldhardy varieties with greater variation in flower color and form. Much of the work in developing cold hardy camellias has been conducted by Dr. Ackerman and Dr. Parks (Aniko 2000). Additionally, longwood Gardens continues a long research program in breeding and selecting camellias (Aniko 2000). The evaluation of woody landscape plants is a long-term commitment, one that often spans the tenures of staff at institutions that Camellias 29 T New Camellia Introductions December 1995 and is 16 feet (4.9 meters) tall and 9 feet (2.7 meters) wide. The single flowers are scarlet red, typical of the species. It received an overall \"A\" ranking, flowered every year, and had especially high marks for foliage quality and habit. This was ranked the number one overall plant of the entire evaluation. It is fully branched to the ground with an excellent ovate habit. `Morris Mercury' (86-050*Z9 \/ KNW 352). One of a group of plants growing on the north side of Gates hall, the Arboretum's administrative offices. This is a precocious, fall blooming plant. This plant has been growing in its current location since October 1999 and is 11 feet (3.4 meters) tall and 7 feet (2.1 meters) wide. It has a more open habit than the others, with an upright arching branch habit. This plant blooms regularly in November of each year, with sporadic blooms the following spring. Despite flowering every year, it received an overall \"B\" rating due to its open habit and foliar damage after cold winters of 2000 and 2001. ANThONY S. AIEllO here are four plants that we are planning to name and introduce. The varietal names and descriptions of these are as follows. All heights are approximate. `Balustrade' (86-043*J \/ KNW 342). One of two plants at the Studio Building, a small office building near our administrative offices. This plant has a very narrow, upright habit and strongly upright branch angles. This plant has been growing in its current location since the spring of 1988 and is 11 feet (3.4 meters) tall and 3 feet (.9 meter) wide. The single flowers are a good scarlet red, typical of the species. It received an overall \"A\" ranking and flowered every year, with excellent lustrous foliage. `Meadowbrook' (86-050*U \/ KNW 352). One of a grove of plants growing on the north side of a large Chamaecyparis pisifera along Meadowbrook Avenue, near the Arboretum's parking lots. This plant has outstanding bluegreen foliage. It has been growing in its current location since December 1995 and is 12 feet (3.6 meters) tall and 6 feet (1.8 meters) wide. Its flower color is a rosy red and lighter in color than others that we have evaluated. It received an overall \"A\" ranking, flowered every year, and had especially high marks for foliage quality; its attractive lustrous foliage stands out for its high quality in all seasons. It is fully branched to the ground with an ovate habit. `Bloomfield' (86-050*W \/ KNW 352). Another in a grove of plants growing on the north side of a large Chamaecyparis pisifera along Meadowbrook Avenue, near the Arboretum's parking lots. This plant combines the best flowering of all of our plants with excellent foliage quality and vigorous growth. This plant has been growing in its current location since Camellia japonica plants growing along Meadowbrook Avenue at the Morris Arboretum. `Bloomfield' (Morris Arboretum 86-050*W ) is pictured at the center of this photograph. 30 Arnoldia 67\/1 Chinese origin (Aiello 2005). likewise, after more than 20 years of evaluation, the Korean Camellia japonica plants represent some of the most cold-hardy collections ever made of common camellia. These collections may extend the hardiness of Camellia japonica into more northern areas and bring the spring pleasure of camellias to eager gardening audiences. Acknowledgements The author would like to thank Elinor I. Goff, Michelle Conners, Shelley Dillard, and Sara ranck for their contributions to this project and to this article. Literature Cited Ackerman, W.l. 2000. Northern Exposure. American Nurseryman 192 (11): 3843. Ackerman, W.l. 2007. Beyond the Camellia Belt. Ball Publishing, Batavia, Illinois. Ackerman, W.l. and D.r. Egolf. 1992. `Winter's Charm', `Winter's hope', and `Winter's Star' Camellias. HortScience 27: 855856. Aiello, A.S. 2005. Evaluating Cornus kousa cold hardiness. American Nurseryman 201 (5): 3239. Aiello, A.S., S. Dillard, E.I. Goff, and M. Conners. 2008. Evaluation of cold hardiness and ornamental characteristics of Korean provenances of Camellia japonica. royal horticultural Society: The rhododendron, Camellia & Magnolia Group Yearbook: 2631. Aniko, T. 2000. Collection Service. American Nurseryman 192 (11): 4450. Bensen, S.D., ed. 2000. New Plants for 2001: Shrubs. American Nurseryman 192 (12): 34. Flint, h.l. 1997. Landscape Plants for Eastern North America. 2nd Edition. John Wiley and Sons, New York. Meyer, P.W. 1985. Botanical riches from afar. Morris Arboretum Newsletter 14 (1): 45. Yinger, B. 1989a. Plant Trek: In pursuit of a hardy camellia. Flower and Garden 33 (2): 104106. Yinger, B. 1989b. Plant Trek: On site with hardy camellias, Sochong Island, Korea. Flower and Garden 33 (3): 6266. Anthony S. Aiello is the Gayle E. Maloney Director of horticulture and Curator at the Morris Arboretum of the University of Pennsylvania in Philadelphia, Pennsylvania. PAUl W. MEYEr A Camellia japonica grown as an espalier in a protected spot at the Morris Arboretum. collect, propagate, and evaluate these plants. At the Morris Arboretum we have found that plants collected in the 1980s in South Korea have exceptional cold hardiness and adaptability. For example, stems from Cornus kousa that were also collected on the 1984 KNW expedition showed significantly more freezing tolerance in tests than plants of either Japanese or "},{"has_event_date":0,"type":"arnoldia","title":"Searching for Exotic Beetles","article_sequence":4,"start_page":31,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25464","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15ea728.jpg","volume":67,"issue_number":1,"year":2009,"series":null,"season":null,"authors":"Campbell, Nichole K.","article_content":"Searching for Exotic Beetles Nichole K. Campbell W hen non-native species of plants, animals, and disease organisms are introduced to other regions they have the potential to become serious pest problems in their new location. Concern over the introduction of potentially damaging species has led the Plant Protection and Quarantine (PPQ) program--part of the USDA Animal and Plant Health Inspection Service--to increase its domestic surveillance for non-native species in the United States over the past several years. Most exotic (non-native) species enter the United States through international movement of people, commodities, and conveyances. Most are accidental introductions, though some intentional introductions (primarily plants) have turned out to be invasive pests. Not all introduced species become agricultural or forest pests; typically, one in seven exotic species is considered invasive. There is often a lapse between the time the pest is introduced and the time that the pest is discovered or reported in the United States; unfortunately this often allows new pest populations to build. Beetle Patrol In 2009, PPQ is conducting exotic beetle trapping around the Boston port area as part of the USDA's national wood borer and bark beetle survey. The Boston port area may be a high-risk PeTer Del TreDICI A cargo ship heads toward the port of Boston, passing between Spectacle Island and Deer Island in Boston Harbor. 32 Arnoldia 67\/1 NICHOle K. CAMPBell area for the introduction of new exotic forest pests because of the high volume of cargo imports that enter the United States through it. Commodities entering the port are often shipped in solid wood packing material, a potential harbor for insect pests of trees. Prior to 2005, there were no regulations requiring the treatment of solid wood packing material for the prevention of pest introductions. Today, all foreign solid wood packing material must be fumigated or heat treated to prevent new forest pests from entering the United States through that very high-risk pathway. The goals of the USDA's national wood borer and bark beetle survey are to obtain information about: The presence, distribution, or absence of target species. The advent of new species. Patterns of distribution throughout the United States and possible pathways for introduction. The phenology of target exotic species in the United States and their selection of hosts. The characteristics of high-risk habitats or sites. The survey methods themselves. When selecting survey sites, we primarily target cargo transport companies, businesses that receive imports, and areas around the port of entry where there are host trees that could support the establishment of exotic beetles. PPQ has chosen twenty locations within 15 miles of the port of Boston for the wood borer and bark beetle survey. One of the sites chosen this year is the Arnold Arboretum because of its close proximity to the Boston port and the presence of a wide variety of tree species in its collections. One of the Lindgren funnel traps at the Arnold Arboretum. Setting the Trap The survey involves trapping and identifying beetles in order to determine if exotic species are present in the area. We placed three lindgren 12-funnel traps at each of the twenty selected locations for a total of sixty traps in the Boston area. each trap is baited with one, or a combina- tion, of the following lures: ultra high release ethanol, ultra high release alpha-pinene, or the 3-ips lure. The volatiles in the lures simulate stressed or dying hardwood and softwood trees, the types of host trees that many of the exotic beetles are attracted to. The traps are hung in trees, on poles, or on fences near target hosts. Traps are placed a minimum of 25 meters (82 feet) apart to prevent volatiles from mixing in the air and deterring beetles. each trap has a collection cup at the bottom that is filled with non-toxic antifreeze to preserve the collected beetles. The trapping period will last from mid March through the end of August to cover a range of emergence periods of the target beetles. Bark and ambrosia beetles typically emerge in early to late spring, while larger wood-boring beetles typically emerge later in summer through fall. The traps are serviced on a bi-weekly schedule to collect any trap contents and replace lures as needed. All of the trapped beetles will be sent to the Carnegie Museum of Natural History, Section of Invertebrate Zoology, in Pittsburgh, Pennsylvania. They will be screened by qualified experts to determine if they are the target exotic beetles or other non-native beetles. Determining the potential invasiveness of these exotic beetles is difficult since there is very little research information available for most of them. Often, they are not studied in their native countries if they do not cause eco- Exotic Beetles 33 nomic damage there. We can't predict exactly how an introduced beetle species will affect forests in the United States, but experts do try to make educated guesses. If any exotic beetles are found they will be confirmed by PPQ experts, and state and local authorities will be notified. The USDA's New Pest Advisory group (part of PPQ), in conjunction with state and local officials, would then evaluate the new pest risk and determine the appropriate action to take to protect our national forests and agricultural industries. A Gallery of Beetles Here are some of the exotic beetles targeted in the survey: WIllIAM M. CIeSlA, FOreST HeAlTH MANAgeMeNT INTerNATIONAl, BUgWOOD.Org Hylurgus ligniperda (Red-haired Pine Bark Beetle) NATIve: europe, Mediterranean areas, Africa, and parts of Asia eNTereD U.S.: Introduced near rochester, New York, in 1994. Found in a lindgren funnel trap. Has been found in four counties. HOST: Pinus spp. (pines) preferred. Also, Abies spp. (firs); Larix spp. (larches); Picea spp. (spruces); Pseudotsuga spp. (Douglas-firs) DAMAge: Affects bark, stem, root, trunk, and seedlings. Feed and develop in tunnels beneath the bark. They are know vectors of the root disease fungi Leptographium spp. and Ceratocystis spp. Ips sexdentatus (Six-toothed Bark Beetle) MIlAN ZUBrIK, FOreST reSeArCH INSTITUTe SlOvAKIA, BUgWOOD.Org Red-haired (or goldenhaired) pine bark beetles under the bark of a Monterey pine (Pinus radiata). Six-toothed bark beetles in galleries. NATIve: Mainland Asia and europe eNTereD U.S.: Has been intercepted at ports of entry. Has not been found domestically beyond ports. HOST: Pinus spp. (pines) preferred. Also, Abies spp. (firs); Larix spp. (larches); Picea spp. (spruces); Pseudotsuga spp. (Douglas-firs) DAMAge: Affects inner bark, leaf, stem, and whole plant. Mates, develops, and feeds in tunnels beneath the bark. Mainly attacks stressed or dying trees. It can kill trees of commercial importance. It also introduces blue stain fungi (Ophiostoma spp.) into host trees which hasten the death of tree, discolor wood, and can result in loss of lumber grade and value. 34 Arnoldia 67\/1 U Two Highly Destructive Exotic Beetles nfortunately, many exotic wood-boring beetles are not attracted to traps baited with volatiles and can only be surveyed for visually. This requires trained spotters using binoculars from ground level, or professional tree climbers knowledgeable in insect signs and symptoms. The Asian longhorned beetle (ALB) (Anoplophora glabripennis) is a highly destructive invasive beetle that can only be surveyed for in this manner. There is ongoing research to identify more effective survey methods for this devastating pest. larvae of the Asian longhorned beetle tunnel into the heartwood of live healthy trees, eventually killing their hosts. Favored species are maples, birches, Ohio buckeye, elms, horse chestnut, and willows. AlB, and efforts to eradicate it, have resulted in the loss of thousands of street trees in several states. AlB was detected in Worcester, Massachusetts, in August, 2008, and its potential spread is of great concern in New england. volunteers will be educated to survey for AlB throughout Massachusetts this year. visual surveys and education outreach for AlB will be conducted in all New england states during 2009. For more information about AlB, please visit: http:\/\/www.aphis.usda.gov\/oa\/alb\/alb.html or http:\/\/massnrc.org\/pests\/alb\/ Emerald ash borer (EAB) (Agrilus planipennis) is another highly destructive beetle that has spread in regions of the United States and Canada. eAB attacks ash trees (Fraxinus spp.) and has been moved from its introduction point in Michigan to other states primarily through movement of nursery stock and firewood. We have not detected eAB in Massachusetts yet, but a survey for it is planned for this year. The Massachusetts Department of Conservation and recreation, Division of Forestry, will place purple panel sticky traps baited with lures at twenty high-risk locations such as campgrounds, nurseries, and wood processors. Currently, there are no plans to trap inside the Arnold Arboretum for eAB because it is not a high-risk location for the introduction of this pest. For more information about eAB, please visit: www.emeraldashborer.info DAvID CAPPAerT, MICHIgAN STATe UNIverSITY, BUgWOOD.Org KeNNeTH r. lAW, USDA APHIS PPQ, BUgWOOD.Org Exotic Beetles 35 NATIve: europe and Asia eNTereD U.S.: Has been intercepted in traps in Indiana (1995) and Maryland (2002). It is not known to be established in the U.S. HOST: Picea spp. (spruces) preferred. Also, Abies spp. (firs); Larix spp. (larches); Pinus spp. (pines); Pseudotsuga spp. (Douglas-firs) DAMAge: Affects bark, crown, foliage, leaf, stem, and whole plant. Considered one of the most serious pests of spruce in europe. It vectors a blue stain fungus (Ceratocystis polonica) which can also kill the host. It causes major economic losses when it is in outbreak numbers and can cause severe decline in spruce populations within its native range. Males aggregate and colonize a stressed tree by boring into the bark and preparing nuptial chambers. The females are then attracted to the chambers to mate. The females lay eggs in maternal galleries where the larva will develop. They can have multiple generations in a year depending on temperature. Dead spruce trees in Slovakia, killed by European spruce bark beetles. Xyleborus seriatus (No common name; very little is known about this beetle.) NATIve: China, russia, Japan, Korea, Taiwan eNTereD U.S.: Intercepted in lindgren trap in Massachusetts in 2005, the first North American record. This beetle was also trapped in Maine in 2008. HOST: Acer spp. (maples), Aesculus spp. (buckeyes), Alnus spp. (alders), Betula spp. (birches), Cryptomeria spp., Fagus spp. (beeches), Larix spp. (larches), Pinus spp. (pines), Prunus spp. (cherries), Quercus spp. (oaks), Thuja spp. (arborvitae), Tsuga spp. (hemlocks), etc. large possible host range. DAMAge: very little data. Is known to be associated with Ambrosiella fungi. Spores of a symbiotic fungi are carried on their bodies to new galleries. larvae and adults feed on this fungi growing between the bark and sapwood. Thought to be a secondary pest and will not kill healthy trees. Several Xyloborus species are potential survey targets. Xylotrechus hircus (No common name; very little is known about this beetle.) NATIve: Native to eastern russia, China, Korea eNTereD U.S.: Intercepted in lindgren trap in Oregon in 1999; not known to be established. HOST: Betula spp. (birches) DAMAge: No information available. Species damage unknown. Several Xylotrechus species are potential survey targets. Nichole K. Campbell is a pest survey specialist with the USDA, APHIS, Plant Protection and Quarantine program. MIlAN ZUBrIK, FOreST reSeArCH INSTITUTe SlOvAKIA, BUgWOOD.Org Ips typographus (European Spruce Bark Beetle) "},{"has_event_date":0,"type":"arnoldia","title":"Early Bloomer: Hydrangea paniculata 'Praecox'","article_sequence":5,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25462","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14e896b.jpg","volume":67,"issue_number":1,"year":2009,"series":null,"season":null,"authors":"Pfeiffer, Sue A.","article_content":"Early Bloomer: Hydrangea paniculata `Praecox' Sue A. Pfeiffer D uring the second half of the nineteenth century, the latest trend in the gardening world was the acquisition and display of exotic plants. At the time, Darwin's theory of evolution was changing the scientific community, and Harvard botanist Asa Gray's paper noting the similarities between eastern North American and eastern Asian floras had recently been published. By the early 1890s, the still young Arnold Arboretum was beginning to take shape. C. S. Sargent, the first director of the Arboretum, had become highly interested in Gray's work comparing our native flora to that of eastern Asia. His desire to plant the Arboretum with every tree capable of surviving the New England climate led him to seek exotic Asian species from similar climates. Although European plant species were easily obtained, acquiring plant material directly from Asia was still difficult during this era. Wanting to view the native flora and personally judge the plants for their landscape value, Sargent set off on a ten week expedition to Japan in the fall of 1892. He collected extensively on the islands of Hondo and Yezo (now known as Honshu and Hokkaido), returning with seeds of 200 species, including Hydrangea paniculata, panicle hydrangea. Hydrangea paniculata is native to Japan and southern Sakhalin Island in Russia as well as eastern and southern China where it is typically found in mixed forests or open hillsides. A large shrub or small tree, panicle hydrangea may reach 20 feet (6 meters) in height, though in New England landscapes a mature height of 10 to 13 feet (3 to 4 meters) is typical. Its large, simple, dark green leaves have toothed margins and a slightly undulating surface. Panicle hydrangea produces conical compound inflorescences 6 to 8 inches (15 to 20 centimeters) in length at the tips of branches. The inflorescences are comprised of two types of florets; a large number of small, cream-colored, fertile florets, plus a scattering of larger, showier, white, sterile florets. The sterile florets often become speckled or flushed with pink as they age. In New England the species flowers from early August into September. When plants were grown from the Hydrangea paniculata seeds collected by Sargent, one was observed to flower far earlier in the summer than the others. Sargent noted this early bloomer in an issue of the Arboretum publication Garden and Forest in September 1897, less than five years after the seed was collected. Several years later, the plant was named `Praecox' (meaning \"premature\") by Arboretum taxonomist Alfred Rehder. Hydrangea paniculata `Praecox' is a vigorous, fast growing, erect shrub which tends to flower three to six weeks earlier than the species. At the Arboretum it typically starts to bloom in early to mid July. Its beauty in the landscape was described in 1922 by Sargent himself: \"When in flower in early July it is one of the handsomest shrubs in the Arboretum,\" and in 1927 by E. H. Wilson: \"Well worth the attention of all interested in hardy plants.\" At the Arboretum, the original plant--now 116 years old--can be found in the Bradley Rosaceous Collection. Although not a member of the rose family, the plant (accession 14714-A) has remained in its original location (formerly the Shrub Collection) because of its importance as a type specimen. The plant is now 15.5 feet (4.7 meters) tall and 24.5 feet (7.5 meters) wide. Every July, visitors are drawn to its incredible display of flowers borne on the many arching stems clad in handsome gray-brown exfoliating bark. Another specimen (accession 14714-1-A), propagated from the original plant in 1905, stands nearby and is equally impressive. Panicle hydrangeas have become very popular in the nursery trade in recent years, and many new cultivars have been introduced. `Praecox' remains the earliest blooming cultivar and is valuable for extending the panicle hydrangea bloom season. While not as readily available as some cultivars, `Praecox' is well worth seeking out and acquiring. Sue Pfeiffer is a Curatorial Fellow at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Index to Volume 66","article_sequence":6,"start_page":36,"end_page":43,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25463","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed15ea36f.jpg","volume":67,"issue_number":1,"year":2009,"series":null,"season":null,"authors":null,"article_content":"Index to Arnoldia Volume 66 Items in boldface refer to illustrations. American cultural\/horticultural studies 2: 2831, 3235 American Society for Horticultural Science 2: 31 Amur corktree, fruit of 1: 14 \"An African Fir Grows in Boston,\" Kyle Port 3: 32, inside back cover Anagnostakis, Sandra L., \"American Chestnuts in the 21st Century\" 4: 2231 Ananas, fruit 2: 25 Andersen, Phyllis, \"A Matter of Taste: Pleasure Gardens and Civic Life\" 3: 1014 Andropogon gerardii 2: 35 Angelo, Ray 3: 5 Anoplophora glabripennis 1: 22 Anthracnose fungus, on ash 4: 35 Apios americana 3: inside front cover Apiosporina morbosa 4: 20, 20 Apples or crabapples with ornamental fruit 2: 22, 23 Arboriculture, for old trees 1: 36 -- 19th-century photo of 2: 29 -- urban 2: 910 Aril 2: 25 Arnold, James 1: 10 Arnold Arboretum, Abies marocana at 3: 32, inside back cover -- -- Accessions process 1: 1718 -- -- -- terms 1: 2021 -- -- Acer saccharinum at 1: 36, inside back cover -- -- African plants in 3: 32 -- -- autumn-fruiting plants 2: 2227, 2227 -- -- Biogeographic Collections 1: 15 -- -- Bussey Hill, in 1930 1: 11 -- -- Case Estates 1: 10, 13 -- -- Cathaya argyrophylla at 3: 2223, 23 -- -- Central Woods 4: 36 -- -- Clethra barbinervis at 4: 36, inside back cover -- -- Conifer Collection 1: 17; 3: 32 -- -- -- -- monotypic rarities in 3: 20, 2223, 23 -- -- conservation collections 1: 16, 17 -- -- Core Collections 1: 1516 -- -- cultivar collections 1: 1718 -- -- Dana Greenhouse and Nursery 1: 19; 3: 22, 32 -- -- Display collections 1: 19 A Abies 3: 16 -- pinsapo var. marocana 3: 32, inside back cover -- -- var. pinsapo `Glauca' 3: 32 Acer 1: 6 -- seeds 2: 26 -- campestre 3: 32 -- dasycarpum 1: 36 -- mono, with bark fungi 4: 18 -- monspessulanum 3: 32 -- negundo, lifespan 2: 2 -- opulus ssp. hispanicum 3: 32 -- saccharinum 1: 36, inside back cover -- -- lifespan 2: 2 -- saccharum 2: 5, 7 -- triflorum, seed 2: 26 \"A Closer Look at Fungi in the Arnold Arboretum,\" Kathryn Richardson 4: front cover, 1321, 13, 1521 Acorn, of bur oak 2: 26 Actinidia spp. 2: 22 -- arguta 4: 36 Adelges tsugae, at Arboretum 1: 2228 Aesculus glabra 2: 36 Africa, plants of 3: 32, inside back cover Agaricus bisporus 4: 14 Aggregate fruits 2: 25 Agricultural record-keeping 3: 5 -- terracing, in China 2: 13 Agrilus planipennis 1: 22 Ailanthus altissima f. erythrocarpa, seeds 2: 26 Albizia julibrissin 1: 6; 2: 27 Alexander, John H. III, \"Collecting Sweetgum in the Wilds of Missouri\" 2: 36, inside back cover Alliaria petiolata 1: 23 \"A Matter of Taste: Pleasure Gardens and Civic Life,\" Phyllis Andersen 3: 1014, 1014 America, the Beautiful (song) 2: 31 \"American Chestnut: The Life, Death, and Rebirth of a Perfect Tree,\" by Susan Freinkel, reviewed 4: 3233 \"American Chestnuts in the 21st Century,\" Sandra L. Anagnostakis 4: 2231, 2230 -- -- documentation and nomenclature 1: 10, 13, 14, 17, 18 -- -- early leadership, and accessions 1: 1013, 1718 -- -- fall color in 2008 4: 35 -- -- flooding in 2008 4: 35 -- -- fungi at 4: 1321, 35 -- -- Hemlock Hill conditions 1: 2228, 2224, 27 -- -- -- -- in 1920 1: back cover -- -- Historic Collections 1: 1718 -- -- Japanese and Korean plants at 4: 36 -- -- introductions by 1: 17, 18 -- -- Larz Anderson Bonsai Collection 1: 13, 18 -- -- Leventritt Shrub and Vine Garden 1: 18 -- -- Liquidambar styraciflua at 2: 36, inside back cover -- -- Living Collections, description and policy 1: 1021 -- -- Malus collection at Arboretum 1: 17, 17 -- -- Meadow Road 1: 21, 36, inside back cover -- -- microclimate project at 4: 35 -- -- Missions statement, and collections 1: 14 -- -- natural areas in 1: 13, 19 -- -- New England flora and 1: 19 -- -- non-native organisms and 1: 2223 -- -- North American taxa 1: 1516 -- -- nursery and horticultural accessions of 1: 17 -- -- Parrotia subaequalis at 1: 9 -- -- Peters Hill 1: 17 -- -- Plant Health Manager 1: 22, 27 -- -- Putnam Fellow 4: 35 -- -- Ralph E. Perry Wood Collection 4: 3 -- -- Rhododendrons at 1: inside front cover, 11, 17 -- -- school children at 1: 28 -- -- seed exchanges 3: 2223 -- -- Spontaneous Flora 1: 14, 19 -- -- Synoptic Collections 1: 16 -- -- tallest tree 1: 36, inside back cover -- -- taxa, botanical and horticultural 1: 10, 13, 17, 18 -- -- Tsuga canadendis, and HWA 1: 2224, 25, 2627 -- -- two-sites issue 1: 10, 13 Index 37 -- -- Weather Station Data--2008 4: 3435 -- -- Willow Path flooding 4: 35 Arnoldia, Index to Volume 65 1: 2935 Aronia melanocarpa, fruit 2: 22, 23 Ascomycota 4: 14 Ash, seeds 2: 26 -- green 2: 5, 6 Ashton, Peter, and collections policies 1: 13 Asia, plants of 1: 29, 10, 12, 15, 27; 2: 1121; 3: 1525; 2630; 4: 2, 6, 710, 36 Asimina triloba 2: 36 Australian Bicentennial Arboretum 3: 17, 18 Autumn fruiting display 2: 2227 Azalea, swamp 1: inside front cover B \"Bai guo\" legend 3: 27, 30 Bailey, Liberty Hyde 2: 29 Bark interest 1: 2, 7, 8, 36; 2: 36; 4: 36 Basidiomycota fungi 4: 13, 14 Bates, Katherine 2: 31 Beach tomato 2: 25 Beautyberry, purple 2: 22, 24 Beech, Japanese 1: 16 Beetle, Asian lady 1: 25 -- -- long-horned 1: 22 -- elm bark 1: 25 -- native lady 1: 25 Berry 2: 22 Betula spp. 2: 7 -- lenta 1: 23, 26; 3: 3, 8; 4: 35 -- nigra 2: 36 -- papyrifera, lifespan 2: 2 -- populifolia 3: 4, 8 Biodiversity, threats to 1: 22 Biogeographic collections 1: 15 Biological responses to climate change 3: 29 Birch, bare-root planting of 2: 7 -- sweet 1: 23, 26, 28; 3: 3, 8; 4: 35 -- river 2: 36 Birds, and fall-fruiting plants 2: 22 Bird's-nest fungi 4: 19 Black knot fungus 4: 20 Blazing star 2: 35 Bloodroot 3: 8 Blueberry, highbush 3: 6, 8 Bluestem, big 2: 35 Blue Hill Observatory, long-term temperatures at 3: 6 Book reviews 2: 2831; 4: 3233 Borer, emerald ash 1: 22 Boston ivy 2: 22 -- parks 3: 10, 13, 14 Boston Public Garden 3: 10, 13, 14 Botanizing in Concord, MA 3: 29 Botany, of fruits 2: 2227 Britton's violet 3: 3 Brooks, Wm. Penn 4: 36 Buartnuts 4: 610 Buckeye, Ohio 2: 36 Buckthorn, glossy 1: 26 Bull, Ephraim 2: 29 Burnham, Charles 4: 25 Bussey Institution 3: 20 Butternut, American, status of 4: 212 -- -- bark 4: 23, 9 -- -- canker disease 4: 46, 4 -- -- cold-hardiness 4: 2 -- -- conservation and restoration 4: 56, 910 -- -- dye from 4: 3 -- -- flowering 4: back cover -- -- genetics 4: 7, chart 8, 910 -- -- native range 4: map 3 -- -- nuts 4: 3, 3, 68, 7 -- -- propagation 4: 6 -- -- regeneration and reproduction 4: 56, 910 -- -- seeds 4: 6, 910 -- -- trees 4: 2, 5, 5, 6, 10 -- -- uses for 4: 24 -- -- wood 4: 34, 3, 4 Buttonbush 2: 36 Buxus 1: 6 C Cactoblastis cactorum 1: 25 Calendar of natural events 3: 5 Callicarpa 2: 22 -- dichotoma, fruiting 2: 24 Canada, butternut status in 4: 4 Capsule 2: 27 Cardinal flower 3: 7 Carnivores, and seed dispersal 2: 18 Carpinus 1: 6 -- caroliniana 2: 36 Carya spp. 4: 2 -- illinoinensis 4: 2 Case Estates 1: 10, 13 Castanea 1: 6 -- crenata 4: 28, 29 -- dentata, blight, and responses to 4: 2233, 2330 -- -- breeding 4: 2829 -- -- decline and regrowth 4: 22, 24, 2829, 3233 -- -- distribution 4: 22, map 22 -- -- flowers of 4: 25 -- -- future of 4: 2231, 32, 33 -- -- gall-wasp damage 4: 29 -- -- genetic analysis of hybrids 4: chart 27 -- -- nuts 4: 27, 28 -- -- regeneration and reproduction 4: 2225, 27, 28, 3233 -- -- trees 4: 23, 26, 27, 30 -- -- wood and timber 4: 22, 2425, 29, 32, 33 -- henryii 4: 29 -- ozarkensis 4: 29 -- pumila 4: 29 -- mollissima 4: 28 Castilleja sessiliflora 2: 35 Castor River [MO] 2: 36 Cathaya, as \"living fossil\" 3: 16, 20 -- argyrophylla 3: 1525, 15, 1819, 21, 2324 -- -- at Arboretum 3: 2223 -- -- collecting in China 3: 1618, 2425 -- -- cultivation in landscape 3: 19, 21 -- -- distribution in China 3: map and chart 1617 -- -- germination record 3: 22 -- -- leaf detail 3: 15 -- -- propagation 3: 18, 18, 2223, 23 -- nanchuanensis 3: 16, 17 \"Cathaya Comes to the Arnold Arboretum\" 3: 2223 \"Cathay Silver Fir: Its Discovery and Journey Out of China,\" Christopher B. Callaghan, with contributions by William McNamara and Peter Del Tredici 3: 1525, 1521, 2324 Cedrus atlantica 3: 32 Celtis occidentalis 2: 5 Central Park, and public taste 3: 10, 11, 14 Cephalanthus occidentalis 2: 36 Cercidiphyllum 2: 17 -- japonicum 4: 36 Channing, William Ellery 3: 5 Cheng, W.C. 3: 20 Cherries 2: 24 Chestnut, American, status and survival 4: 2231; 3233 -- -- history and culture 4: 22, 2829, 3233 -- Asian species 4: 2324, 28 38 Arnoldia 67\/1 -- breeding 4: 2327, 2831, 32 -- hybrids 4: chart 27 -- nut-growing 4: 23, 24, 27, 2829, 28 Chestnut blight disease 4: 14, 2231, 24, 3233 -- -- -- biological controls 4: 2426, 27, 28 -- -- -- resistance to 4: 2425, 26, 28 -- -- gall wasp 4: 29, 29 -- -- weevil 4: 28, 29 Chicken of the woods fungi 4: 17 China, nature reserves in 3: 1718 -- plants of, 1: 29, 12, 27; 2: 1121; 3: 1525, 2630 \"Chinese Parrotia: A Sibling Species of the Persian Parrotia,\" Jianhua Li and Peter Del Tredici 1: 29, 28 Chokeberry 2: 23, 23 Christopher B. Callaghan, with contributions by William McNamara and Peter Del Tredici, \"Cathay Silver Fir: Its Discovery and Journey Out of China\" 3: 1525 Chryphonectria parasitica fungus 4: 2223, 3233 -- -- -- biological controls of 4: 2628, 32 -- -- -- in Connecticut, circa 1900 4: map 24 Chung, H.C. 3: 16 Civilian Conservation Corps (CCC), 1930s 2: 34, 34 Civil War uniforms, butternut-dyed 4: 3 Clark, Stacy 4: 29 Clethra alnifolia 2: 27; 4: 36 -- barbinervis 4: 36, inside back cover -- -- centenarian at Arboretum 4: 36 Climate change, biological responses to 3: 29 -- -- ginkgo's adaptability to 2: 17, 18 \"Collecting Sweetgum in the Wilds of Missouri,\" John H. Alexander III 2: 36, inside back cover Clover, prairie bush 2: 35 Cold-hardiness 2: 36; 4: 2 Collections policy 1: 1021 Colorado College 2: 31 Compass plant 2: 35 Compression of tree trunks by roots 2: 47, 810 Concord, MA, flora of and climate change 3: 29 -- -- preservation property in 3: 9 -- -- Public Library archives 3: 5 Coneflower, prairie 2: 35 Conifers 1: 2228; 3: 1525; 4: 17, 20 -- dwarf collection at Arboretum 1: 17 -- fungi on 4: 17, 20 Connecticut Agricultural Experiment Station (CAES) 4: 22, 24, 26, 28, 29 Connecticut, chestnuts in 4: 22, 23, 24 -- -- -- circa 1900 4: 23, 24, 30 -- HWA in 1: 25, 26 Conservation of forestland 3: 9, 32 Convallaria majalis, flowering and winter temperatures 3: 5 Coop, Julie 4: 35 Coprinus comatus 4: 18, 18 Cornus, anthracnose and 4: 35 -- fruiting 2: 24 -- alternifolia, flower 1: 15 -- controversa 1: 8, 15 Corylopsis sinensis var. glandulifera, blossom with bee 4: inside front cover Cotinus coggyria 1: 12 Cotoneaster 2: 23 Crabapple `Donald Wyman' 2: 23 -- urban planting 2: 7 Cranberrybush 2: 24 Crataegus 2: 23 Crucibulum fungi 4: 19 Cucumbertree magnolia, senescent at Arboretum 4: front cover \"Curatorial Notes: An Updated Living Collections Policy at the Arnold Arboretum,\" Michael S. Dosmann 1: 1021, 1112, 1421 Curtis, James 2: 31 Curtis Prairie 2: 3235, 33 Cyathus fungi 4: 19 -- -- -- \"The Li Jiawan Grand Ginkgo King,\" with Zhun Xiang, Yinghai Xiang, and Bixia Xiang 3: 2630, 2730 -- -- -- \"Wake Up and Smell the Ginkgos\" 2: 1121 -- -- -- photos by 2: front\/back covers Denmark, public garden 3: back cover Diospyros 1: 6 -- virginiana, fruit of 2: 22, 22 Disjunct flora 1: 24, 15, 15 Disney, Walt, and public parks 3: 14 Distyliopsis tutcheri 1: 3 Distylium racemosum 1: 3 Dog vomit slime mold 4: 21 Doogue, William 3: 10, 14 Dosmann, Michael S., \"Curatorial Notes: An Updated Living Collections Policy at the Arnold Arboretum\" 1: 1021 -- -- -- photo by 1: inside front cover Drupe 2: 24, 25 Dutch elm disease 1: 25; 4: 14 \"Dysfunctional Root Systems and Brief Landscape Lives: Stem Girdling Roots and the Browning of Our Landscapes,\" Gary Johnson 2: 210, 36, 810 E Ecosystem disturbance 1: 2627 \"Ecosystems in Flux: The Lessons of Hemlock Hill,\" Richard Schulhof 1: 2228, 2224, 27, 28 Edinburgh Conifer Conservation Program 3: 18 Elm, American 1: 25 -- arborists in, 1890s 2: 29 -- seed 2: 26 Elysian Fields park [NJ] 3: 12 Emerald Necklace parks 3: 13 Endangered or rare plants 1: 29, 17, 2228; 2: 1121; 3: 1525, 2630; 4: 212, 2231, 3233 English yews 2: front\/back covers Enkianthus campanulatus 1: front cover Environmental education 3: 9 Ergot fungus 4: 1415 Estabrook Woods 3: 9 Evolution, of Hamamelidaceae 1: 26 \"Excerpt From Fruits and Plains: The Horticultural Transformation of America,\" Philip J. Pauly 2: 3235, 3335 D Daniel Boone National Forest [KY] 4: 2 Dean, Brad 3: 5 Death camas 2: 35 Debreczy, Zsolt and Istvan Racz, photo by 1: inside back cover Deer damage to trees 4: 5, 29 Delaware Valley, Clethra in 4: 36 Delphinium carolinianum subsp. virescens 2: 35 Del Tredici, Peter, \"Cathay Silver Fir: Its Discovery and Journey Out of China,\" with Christopher B. Callaghan, and William McNamara 3: 1525 -- -- -- \"The Chinese Parrotia: A Sibling Species of the Persian Parrotia,\" and Jianhua Li 1: 29 Index 39 F Fabaceae, seeds of 2: 27 Fagus 1: 6 -- crenata 1: 16 Fairchild, David 2: 31 Fairchild Botanical Garden 2: 31 Fairy Lake Botanical Garden 3: 19, 22 Famiglietti, Bob 4: 35 Farrand, Beatrix 1: 10 Fassett, Norman, and prairie landscape 2: 3235 Faxon, Charles, illustration by, circa 1900 4: back cover Felis bengalensis 2: 18 Fir, African 3: 32, inside back cover -- Douglas 3: 23 Firethorn 2: 23 Florida, early horticulture in 2: 31 Flowering times and climate change 3: 29 Flycatcher, pied 3: 3 Follicle 2: 25 Forest Hills Cemetery, ginkgos in 2: 15, 16 Forests, Caspian [Central Asia] 1: 6 -- conservation of 3: 9 -- Eastern 4: 56, 10, 22, 28, 32 -- hardwood 4: 5, 22 -- loss of mixed in China 2: 12, 13 Forsythia `Courdijau' 1: 18 Fortunearia fortunei 1: 8 Fothergilla major 1: 3 Fragrant plants 4: 36 Frangula alnus 1: 26 Fraxinus, spp. 2: 7 -- anthracnose on 4: 35 -- seeds 2: 26 -- pennsylvanica 2: 5 Freinkel, Susan, \"American Chestnut: The Life, Death, and Rebirth of a Perfect Tree,\" reviewed 4: 3233 Fruiting, botany of autumn 2: 2227 \"Fruits and Plains: The Horticultural Transformation of America,\" Philip J. Pauly, reviewed and excerpted 2: 2831, 3235 \"Fruits of Autumn,\" Nancy Rose 2: 2227, 2227 Fu, Chengxin 1: 6 Fuligo septica 4: 21, 21 Fungi, at Arboretum 4: front cover, 1321 -- classification and identification of 4: 1315 -- hardwoods and 4: 15, 17, 18 Fungus disease, of butternut 4: 211 -- -- of chestnut 4: 14, 2231, 3233 -- -- of elm 1: 25; 4: 14 G Galloway, Beverly T. 2: 29 Ganoderma lucidum 4: 21 -- tsugae 4: 20, 20 Gardens, public 1: 1021; 3: 1014 Garlic mustard 1: 23 Geranium `Crystal Palace Gem' 3: 10, 10 Ghizhou Botanical Garden 3: 17 Ginkgo, ancient giant of Li Jiawan, China 3: 2630, 27, 29 -- -- -- drawings of, in elevation and cross-section 3: 28, 30 -- -- -- folk legends 3: 27, 30 Ginkgo adiantoides, in fossil record 2: 17 -- biloba 2: 1121, 11, 12, 1415; 3: 2630 -- -- adaptability 2: 11, 13, 17, 18, 20 -- -- aging and lifespan 3: 2627, 30 -- -- epiphytes on 2: 12 -- -- evolutionary history 2: 1718 -- -- foliage: 2: 11, 11, 13, 17, 19 -- -- fruits 2: 1516, 15, 18 -- -- -- dispersal by animals 2: 18 -- -- genetic diversity 2: 12 -- -- growth 2: 14, 14, 18; 3: 2627, 28, 29, 30, 30 -- -- in cultivation and the wild 2: 1214, chart 14, 17 -- -- pollination 2: 15 -- -- popularity 2: 11, 12, 13; 3: 27, 30 -- -- populations 2: 1121 -- -- regeneration 3: 2627, 30 -- -- reproduction 2: 1318 -- -- -- and latitude 2: chart 16 -- -- -- and temperature 2: 15, 1617 -- -- urban planting 2: 11, 11, 18, 19, 20 -- yimaensis, in fossil record 2: 17, 17 Gleditsia spp. 2: 27 -- triacanthos, pods of 2: 27, 27 -- -- `Shademaster' 2: 5 Global climate change 3: 2, 3, 6 Glyptostrobus 2: 17 Golden rain tree capsules 2: 27, 27 Goldenrods and climate change 3: 8 Gong, Wei 2: 13 Grape 2: 22 -- honeysuckle 2: 22 Graves, Arthur 4: 24 Gray, Asa 1: 3 Great Meadows National Wildlife Refuge 3: 9 Great Smoky Mts. National Park, HWA in 1: 26 Greenbrier 2: 35 Grey Towers historic site, butternut interior 4: 4, 4 Grifola frondosa 4: 19, 19 Groundnut 3: inside front cover Guangxi Institute of Botany 3: 16 Gustafson, Kathryn, and Crosby, Schlessinger, and Smallwood (designers) 3: 14 Gymnocladus dioicus 2: 27 Gypsy moth removal, 19th-century photo 2: 29 H Hackberry 2: 5, 6 Hallucinogen in grain fungus 4: 1415 Hamamelidaceae, evolutionary kinships 1: 26, chart 3 -- genetic analysis 1: 34 Hamamelis 1: 24 -- x intermedia `Jelena' 3: 31 -- japonica 1: 2 -- mexicana 1: 2 -- mollis 1: 2 -- vernalis 1: 2 -- virginiana 1: 2, 3 Hardwood Tree Improvement and Regeneration Center 4: 10 Hardwood forests 4: 5 -- fungi 4: 15, 17, 18 Harmonia axyridis 1: 25 Harvard Forest, HWA study 1: 27 Hawthorne 2: 23 He, Shan-an 1: 6 Heartnuts 4: 7, 7 Heimarck, Heather D., book review by 4: 3233 Helianthus rigidus 2: 35 Hemlock, Chinese, and HWA 1: 27 -- eastern 1: 2228 Hemlock varnish shelf fungus 4: 20 -- woolly adelgid (HWA), at Arboretum 1: 2228 -- -- -- cold and 1: 26 -- -- -- egg masses of 1: 22 -- -- -- history of 1: 25, 26 -- -- -- management of 1: 2228 -- -- -- pesticides for 1: 26 -- -- -- resistance to 1: 27 Hen of the woods fungi 4: 19 Hessian fly 2: 29 40 Arnoldia 67\/1 Hickories 4: 2 Hird, Abby 4: 35 Hokkaido, plants of 4: 36 Hollies 2: 22, 24, 24 Honey locust 2: 5, 6, 27, 27 Honeysuckle berries 2: 22, 22 Hornbeam, American 2: 36 Horticultural oil 1: 26 Horticulture, civic 3: 1014 Hosack, David 2: 29 Hosmer, Alfred 3: 29 Hovey, Charles 2: 29 Hsu, H.L. 3: 16 Huanyong, Chen 3: 16, 20, 20 Huaping Nature Reserve 3: 17 Hummingbirds and climate change 3: 9 Hunnewell, Horatio Holis 2: 31 Hunnewell Pinetum, 1901 image 2: 30 Hurricane damage to maples 1: 36 -- of 1938 1: 25 Hybrids, role in species recovery 4: 9 Hydrangea paniculata `Praecox' 1: 18 Hypovirus 4: 26, 28 I Ilex spp. 2: 22, 24 -- decidua 2: 36 -- verticillata `Red Sprite' 3: front cover -- -- `Winter Red' 2: 24 Imidacloprid 1: 26 \"Index to Arnoldia, Volume 65\" 1: 2935 Inky cap mushrooms 4: 18, 18 International Registrar for Chestnut Cultivars 4: 29 International Union for Conservation of Nature and Natural Resources 3: 32 Introduced organisms 1: 2223, 25, 2627 Ironwood, Chinese 1: 29 -- Persian 1: 29 Jefferson, Thomas, and horticulture 2: 29 -- -- recordkeeping 3: 5 Jeholornis, and ginkgos 2: 18 Jensen, Jens 2: 29, 32 Jiangsu Institute of Botany 1: 4 Jixin, Zhong 3: 16, 24 Johnson, Gary, \"Dysfunctional Root Systems and Brief Landscape Lives: Stem Girdling Roots and the Browning of Our Landscapes\" 2: 210 Johnson, Samuel 3: 11 Juglandaceae 4: 2 Juglans 4: 2 -- ailantifolia 4: 2 -- -- var. cordiformis 4: 7 -- x bixbyi 4: 2, 6 -- Cardiocaryon Section 4: 2 -- cinerea, status and survival 4: 212, 27, 9, 10 -- -- bark 4: 2, 3, 4, 6, 9 -- -- canker 4: 46, 4 -- -- `Chamberlin' 4: 3 -- -- `Craxezy' 4: 3 -- -- DNA 4: 910 -- -- fungus epidemic 4: 211 -- -- hybrids compared to species 4: 610, 6, 7, chart 8 -- -- -- nuts 4: 2, 3, 3, 5, 69, 67 -- -- inflorescence, circa 1900 illustration 4: back cover -- -- lenticels 4: 9 -- -- native range 4: map 3 -- -- nuts 4: 3, 7 -- -- trees 4: 26, 2, 5, 6, 10 -- -- wood 4: 3, 3, 4, 4 -- mandshurica 4: 2 -- nigra 4: 2 -- x quadrangulata 4: 2 -- regia 4: 2 -- Rhysocaryon Section 4: 2 -- Trachycaryon Section 4: 2 Koller, Gary, plant-collecting anecdote 2: 36 Korea, plants of 1: 12; 2: 23; 4: 36 Kwangfu-Lingchu Expedition 3: 16 Kwangtung Institute of Botany 3: 16 L Laetiporus cincinnatus 4: 17 -- sulphureus 4: 17, 17 Larkspur, prairie 2: 35 Latinized cultivar names 1: 13, 18 Lei, H.C. 3: 16 Leopard cat 2: 18 Leopold, Aldo, and prairie culture 2: 31, 3235 Lepachys pinnata 2: 35 Lespedeza capitata 2: 35 Li, Jianhua, \"The Chinese Parrotia: A Sibling Species of the Persian Parrotia,\" with Peter Del Tredici 1: 29 \"Li Jiawan Grand Ginkgo King,\" Zhun Xiang, Yinghai Xiang, Bixia Xiang, and Peter Del Tredici 3: 2630, 2730 Liatris spp. 2: 35 Lichens 4: 14 Livestock, plants poisonous to 2: 35 Linden 3: back cover -- littleleaf 2: 58, 56, 8, 9 Liquidambar styraciflua, in Missouri 2: 36 -- -- bark 2: inside back cover Living Collections Policy 1: 1021 -- -- -- text of 1: 1421 \"Living fossils\" 2: 18; 3: 1517, 20 Lobelia cardinalis 3: 7 Longenecker, William 2: 3233, 34 Longwood Gardens 4: 32 Lonicera 2: 22 -- reticulata, fruit 2: 22 Lupine, sundial 2: 35 Lupinus perennis 2: 35 M Maclura pomifera, fruit 2: 25, 25 Magnolia, fruiting behavior 2: 25 -- sweetbay hybrid 2: 25, 25 Magnolia acuminata, senescent 4: front cover -- virginiana, seeds 2: 25, 25 Malus spp. 2: 7, 22, 23 -- -- collection at Arboretum 1: 17, 17 -- `Donald Wyman' 2: 22, 23 J Jack, John 1: 17; 3: 20, 20 Japan, plants of 1: 18; 4: 2, 6, 7, 36 \"Japanese Clethra: A Hidden Gem,\" Richard Schulhof 4: 36, inside back cover Japanese clethra, centenarian specimen 4: 36, inside back cover -- knotweed 1: 23, 26 K Katsura 4: 36 Keeteleria fortunei 3: 16 Kentucky coffee tree 2: 27 Keren, Kuang 3: 16 Kiwi, hardy 2: 22; 4: 36 Knotweed, Japanese 1: 23, 26 Koelreuteria paniculata capsules 2: 27, 27 Index 41 Maple, Norway, compression of stem tissue 2: 5 -- seeds 2: 26, 26 -- silver 1: 36, inside back cover; 2: 2 -- sugar 2: 5, 6, 7, 8 Marlatt, Charles L. 2: 29 Massachusetts, climate change in 3: 29 Massachusetts Gypsy Moth Commission 2: 29 Massachusetts state agricultural college 4: 36 Mayetiola destructor 2: 29 McFarland, John Horace, 1920 photo by 1: back cover McNamara, William 3: 18 -- -- \"Cathay Silver Fir: Its Discovery and Journey Out of China,\" with Christopher B. Callaghan and Peter Del Tredici 3: 1525 -- -- \"Three Conifers South of the Yangtze\" excerpted 3: 2425 Medicinal use of fungi 4: 21 Mendocino Botanic Garden 3: 25 Merkel, Hermann 4: 32 Merrill, Elmer D. 3: 15 Metasequoia 2: 17 -- glyptostroboides 3: 15, 20 Millenium Park, Chicago 3: 14 Miller, Wilhelm 2: 32 Miller-Rushing, Abraham J., \"Impact of Climate Change on the Flora of Thoreau's Concord\" with Richard J. Primack 3: 29 -- -- -- photo by 3: inside front cover Minnesota Department of Forest Resources, urban tree survey 2: 510 Missouri, collecting sweetgum in 2: 36 Missouri Botanical Garden herbarium 4: 9 Moroccan fir 3: 32, inside back cover Morocco, fir forest in 3: 32 -- plants of 3: 32 Morus, fruit 2: 25 Moth, winter 1: 23 Mountain ash, autumn interest 2: 22, 23 -- -- Korean 2: 23, 23 -- habitats 1: 68; 2: 1213; 3: 16, 17, 1921, 26, 32 Mulberry 2: 25 Mulch, mold in 4: 21 Munroe, Alfred, 1890s photo by 3: 4 Murrill, William 4: 32 Mutualist fungi 4: 14 Mycena spp. 4: 1819, 18 Mychorrhizae and trees 4: 14 Myxomycetes 4: 15, 21 N Nanjing Botanical Garden 1: 6, 7 Nannyberry, fruits of 2: inside front cover National Clonal Germplasm Repository [Davis, CA] 4: 7, 9 Native Americans 2: 33; 4: 23, 32 Nature Conservancy 4: 10 New England flora 1: 19 -- -- -- and climate change 3: 29 New York, public gardens in 3: 10, 1112 New York Botanical Garden 3: 22 Nicholson, Rob 3: 32 Nidulariales fungi 4: 19 Nolen, John 2: 32 Non-native organisms 1: 2229; 4: 212; 2233 North America, plants native to 1: 1516, 2228, 36; 2: 2331, 36; 3: 3235; 4: 212, 2231, 3233 North American Plant Collections Consortium (NAPCC) 1: 1516 Northern Research Station of Forest Service [MN] 4: 6 Nurseries, historic 1: 17 Nursery trade, post-World War II 1: 13 Nut trees 2: 1518; 4: 212, 2233 Nyctereutes procynoides 2: 18 O Oak, bur (acorns) 2: 26 Olmsted, Frederick Law, Jr. 2: 29 Olmsted, Frederick Law, Sr. 2: 29, 31; 3: 1011 Olmsted park design 2: 32; 3: 1011 Operaphtera brumata 1: 23 Ophiostoma ulmi 1: 25 Opuntia spp. 1: 25 -- macrorhiza 2: 35 Ostry, Michael 4: 6 Oudolf, Piet 3: 14 Oxalis stricta 3: 6 Oyster mushroom 4: 17, 17 P Paeonia coriaceae var. marocana 3: 32 Paguma larvata 2: 18 Palm civet 2: 18 Parasitic fungi 4: 14, 15, 17, 19 Parks and popular aesthetics 3: 1014 Parrot, F.W. 1: 4 Parrotia, Chinese 1: 29 Parrotia, botanical literature on 1: 23, 45, 6 -- distribution 1: map 2 -- family relationships 1: chart 3 -- persica 1: 26 -- subaequalis 1: 29, 38 -- -- at Arboretum 1: 9 -- -- evolution and genetics 1: 26 -- -- penjing (bonsai) subject 1: 6 -- -- pollination 1: 4 -- -- propagation: 1: 6, 6, 7, 7, 9 -- -- Shaniodendron and 1: 4 Parrotiopsis jacquemontana 1: 3 Parthenocissus 2: 22 Pauly, Philip J., \"Fruits and Plains: The Horticultural Transformation of America,\" title reviewed and excerpted 2: 2831, 3235 -- -- -- death of, in 2008 2: 31 Pawpaw 2: 36 Payne, Jerry 4: 29 Peaches 2: 24 Pear 2: 23 -- Chinese sand 2: 23 Pecan 4: 2 Penjing, Parrotia as 1: 6 Persimmon 2: 22, 22 Pest control, unintended effects of 1: 25, 26 Pheasant's-back polypore 4: front cover, 15, 15 Phenological events and flowering times 3: 29 Phellodendron amurense 1: 14 Picea 3: 23 Picturesque landscape ideals 3: 1011 Pijut, Paula M., \"The Peril and Potential of Butternut,\" with Keith Woeste 4: 212 Pinaceae 3: 23 Pinchot, Gifford, home of 4: 4 Pineapple 2: 25 Pine, jack 2: 2 Pinus spp. 3: 16, 23 -- banksiana, lifespan 2: 2 Planting depth, and lifespan of city trees 2: 4, 510 Platanus 2: 17 Pleasure garden, aesthetics of 3: 1014 42 Arnoldia 67\/1 Pleurotus ostreatus 4: 17, 17 Plums 2: 24 Png, S.K. 3: 17 Pods 2: 27 Pogonia ophioglossoides 3: 7 Poison ivy 2: 35 Pollinators and climate change 3: 3, 89 Polygonum cuspidatum 1: 23 Polyporus squamosus 4: 15, 15 Pome 2: 23 Pomology, in 19th-century America 2: 2831, 3235 Port, Kyle, \"An African Fir Grows in Boston\" 3: 32, inside back cover Possumhaw 2: 36 Prairie landscape 2: 3235, 33 -- -- species and weeds 2: 3435, 35 -- painted cup 2: 35 -- rose 2: 35 Prickly-pear cactus 2: 35 Primack, Richard J., \"The Impact of Climate Change on the Flora of Thoreau's Concord,\" with Abraham J. Miller-Rushing 3: 29 -- -- -- photo by 3: inside front cover Prunus, edible fruit in genus 2: 24 Pseudocolus fusiformis 4: 1617, 16 Pterocarya 1: 6 Pterostyrax corymbosum 1: 8 Public gardens and aesthetics 3: 1014 Pyracantha 2: 23 Pyrus pyrifolia, fruit 2: 23, 23 Rhododendron viscosum f. rhodanthum 1: inside front cover -- yedoense var. poukhanense 1: 11 Richardson, Kathryn, \"A Closer Look at Fungi in the Arnold Arboretum\" 4: front cover, 1321 Riming, Hao 1: 4, 6, 6 Riparian habitats 1: 8; 2: 17, 36 Rock, Joseph 1: 17 Root-growth, of yews 2: front\/back covers Root restriction, and tree mortality 2: 210, 46, 810 Rosa carolina 2: 35 -- rugosa hips 2: 25 Rose, Nancy, \"Silver Wins Gold\" 1: 36, inside back cover -- -- \"The Fruits of Autumn\" 2: 2227 -- -- photos by 1: front cover; 2: inside front\/inside back covers; 3: front\/inside back covers; 4: front\/ inside front covers Rose, prairie 2: 35 Rose family 2: 23 -- pogonia 3: 7 Rosehips 2: 25 Rosinweed 2: 35 Royal Botanic Garden, Edinburgh 3: 18 Royal Botanic Gardens, Sydney 3: 18 Russia, plants of 1: 12 S Salem witch trials, and ergot poisoning 4: 15 Samara 2: 26, 26 Sanguinaria canadensis 3: 8 Sapphireberry 2: 24 Sapporo Agricultural School 4: 36 Saprobic fungi 4: 14, 15, 17, 19 Sargent, Charles S. 1: 10, 17, 18; 2: 29 Saunders, William 2: 29 Sax, Karl, and Peters Hill 1: 17 Schlereth, Thomas J., book review by 2: 2831 Schneider, Stephen 4: 35 Schulhof, Richard, \"Ecosystems in Flux: The Lessons of Hemlock Hill\" 1: 2228 -- -- \"Japanese Clethra: A Hidden Gem\" 4: 36 -- -- photo by 4: inside back cover Seasonal temperatures and flowering times 3: 78 Q Quackgrass 2: 34 Quarryhill Botanic Garden 3: 18 Quercus spp. 1: 6; 2: 26; 4: 35 -- fungi of 4: 17, 19 -- macrocarpa, lifespan 2: 2 -- -- acorns 2: 26 Qiu, Ying-xiong 1: 7 Qiu, Yinlong 1: 4 R Raccoon dog 2: 18 Racz, Istvan, and Zsolt Debreczy, photo by 1: inside back cover Raspberry, fruits 2: 25 Raynal, Guillaume 2: 29 Rhododendron collection at Arboretum 1: 17 Seed dispersal, of ginkgos 2: 18 Shaniodendron 1: 45 Silk-tree 2: 27 Silphium spp. 2: 34, 35 -- laciniatum 2: 35 Silva of North America, illustration from, circa 1900 4: back cover \"Silver Wins Gold,\" Nancy Rose 1: 36, inside back cover Simberloft, Daniel 2: 29 Simpson, Charles T. 2: 29 Sirococcus clavigignenti-juglandacearum, discovery and description 4: 45 Slime mold 4: 21 Smilax lasioneura 2: 35 Soil, excess and tree decline 2: 510 Solidago graminifolia 3: 8 -- rigida 2: 35 -- rugosa 3: 8 Sorbus 1: 6; 2: 22, 23 -- alnifolia 2: 23, 23 -- yuana 1: 12 Sorrel, yellow wood 3: 6 Southwest China, Off the Beaten Track 3: 17 Spain, plants of 3: 32 Sperry, Theodore, and prairie style 2: 32, 3435 Spruces 3: 23 Stewartia rostrata 1: 8 -- sinensis 1: 8 Stinkhorn fungi, in Arboretum 4: 1617 Storm damage, to urban trees 2: 2, 3, 45, 7, 89 Styrax confusus 1: 8 Sukachev (Soviet botanist) 3: 16 Summersweet 2: 27, 27; 4: 36 Sunflower, stiff 2: 35 Sun Yat-sen University 1: 4 Sweetbay magnolia seeds 2: 25 Sweetgum, collecting in Missouri 2: 36, inside back cover Sycamore 2: 36 Sycopsis sinensis 1: 3 Symplocos paniculata, fruit 2: 24 Syringa, at Arboretum 1: 17 T Talassemtane National Park 3: 32 Tan, H.F. 3: 16 Index 43 Taxus 1: 6 -- baccata 2: front\/back covers \"The Impact of Climate Change on the Flora of Thoreau's Concord,\" Abraham J. Miller-Rushing and Richard J. Primack 3: 29, 25, 7, 8 \"The Peril and Potential of Butternut,\" Keith Woeste and Paula M. Pijut 4: 212, 27, 9, 10 Thoreau, Henry David, and flora of Concord, MA 3: 29 -- -- -- statue at Walden 3: 2 \"Three Conifers South of the Yangtze,\" William McNamara, excerpted 3: 2425 Tianping Mts. [China] 3: 16 Tilia cordata, and root-girdling 2: 5, 5, 7, 8, 8, 9 Tivoli Garden 3: 13, 13, back cover Tokyo, gingkos in 2: 11, 20 Toxicodendron radicans 2: 35 Trabut, Louis Charles 3: 32 Trametes versicolor 4: 18, 18 Tree-of-heaven, red-seeded 2: 26 Trees, centenarian or notable specimens 1: 21, 36, inside back cover; 2: front\/back covers, 29; 3: 2630, 27, 29; 4: front\/inside back covers, 15, 30, 36 -- epidemic losses of 1: 2228; 2: 210; 4: 212, 14, 2233 -- relationships with fungi 4: 1321 -- restricted growth, and lifespan 2: 210, chart 2 -- prairie 2: 33, 34 -- urban plantings 2: 210, 9, 11, 20 -- veneration of, in Asia 3: 30 Tremella mesenterica 4: 16, 16 Tsuga canadensis, decline due to HWA 1: 2228 -- -- fungus on 4: 2021, 20 -- chinensis, landscape potential 1: 27 Turkey-foot grass 2: 35 -- tail fungus 4: 18, 18 \"Two Living Fossils and the Arnold Arboretum Connection\" 3: 20 UVW Ulmus seeds 2: 26 -- americana 1: 25 University of California, Berkeley 3: 23 University of Minnesota, urban forestry research 2: 510 University of New Hampshire 1: 4 University of Wisconsin Arboretum 2: 31, 3235, 33 Urban tree losses 2: 210 -- ginkgo plantings 2: 11, 20 -- park displays 3: 1014 USA National Phenology Network 3: 9 USDA Forest Service 4: 4, 10, 29 -- -- -- -- inspections 1: 22 -- -- -- -- -- -- Northern Research Station [MN] 4: 6 US National Academy of Sciences 1: 22 US National Arboretum 2: 31 US National Park Service 2: 34 Vaccinium corymbosum 3: 6 Van Rensselaer, Mariana 3: 10 Vaux, Calvert 3: 10, 11 Vauxhall Gardens [London] 3: 1112, 13 -- -- engraving of, 1785 3: 12 Veblen, Thorstein 3: 11 Vernalization 3: 67 Viburnum 2: 22 -- lentago, fruit 2: inside front cover -- trilobum, fruit 2: 24 Viola brittoniana 3: 3 Virginia, HWA in 1: 25 Virginia creeper 2: 22 Vitis spp. 2: 22 \"Wake Up and Smell the Ginkgos,\" Peter Del Tredici 2: 1121, 1115, 1720 Walden, by Henry David Thoreau 3: 5 Walden Pond 3: 45, 9 -- -- in 1890s 3: 4 Walnut, eastern black 4: 2 -- English 4: 2 -- Japanese, and hybrids of 4: 2, 69, 9 -- Manchurian 4: 2 -- Persian 4: 2 Walnuts, New World 4: 2 Ward, Samuel A. 2: 31 \"Weather Station at Arboretum--2008 Summary\" 4: 3435, 35 Weigela subsessilis 1: 12 Wellesley College, and horticultural heritage 2: 30, 31 Wilson, Ernest Henry 1: 17; 3: 21; 4: 36 Winterberry `Red Sprite' 3: front cover -- `Winter Red' 2: 24 Winter temperatures, and flowering times 3: 67, 8 Wisconsin, butternut canker in 4: 4 -- prairies 2: 3235, 33, 34 Wisteria spp. 2: 27 Witchhazels 1: 25 Witch's butter fungus 4: 16 -- -- -- European folk legends of 4: 16 Witch trials and ergot poisoning 4: 1415 Woeste, Keith, \"The Peril and Potential of Butternut,\" with Paula M. Pijut 4: 212 Wollemia nobilis 3: 15 Wollemi pine 3: 15 Wood, compression by girdling roots 2: 47, 810 -- tissue, healthy and malformed 2: 5 Woodland Period Indians, in Wisconsin 2: 33 Woody plants, fruits of 2: 2227, 2227 Wyman, Donald, and collections policies 1: 10, 13, 17 -- -- eponymous crabapple 2: 23 XYZ Xianfu, Deng 3: 16 Xiang, Bixia, \"The Li Jiawan Grand Ginkgo King,\" with Zhun Xiang, Yinghai Xiang, and Peter Del Tredici 3: 2630 Xiang, Yinghai, \"The Li Jiawan Grand Ginkgo King,\" with Zhun Xiang, Bixia Xiang, and Peter Del Tredici 3: 2630 Xiang, Zhun, \"The Li Jiawan Grand Ginkgo King,\" with Yinghai Xiang, Bixia Xiang, and Peter Del Tredici 3: 2630 Xiansu, Hu 3: 20 Xu, Shixian 3: 27, 28 Yew, English 2: front\/back covers Yin, Dr. (Chinese botanist) 3: 24 \"Yinshan\" 3: 24 Yixing Caves 1: 7 Yong, Li 3: 22 Zelkova 1: 6 Zhejiang University 1: 7; 2: 13 Zhong, Jixin 3: 24 Zigadenus elegans 2: 35 Index compiled by Rosalie Davis. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23414","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160a728.jpg","title":"2009-67-1","volume":67,"issue_number":1,"year":2009,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Peril and Potential of Butternut","article_sequence":1,"start_page":2,"end_page":12,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25459","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14e8528.jpg","volume":66,"issue_number":4,"year":2009,"series":null,"season":null,"authors":"Woeste, Keith; Pijut, Paula M.","article_content":"The Peril and Potential of Butternut Keith Woeste and Paula M. Pijut AMy ROSS-DAvIS B utternut (Juglans cinerea), also known as white walnut because of its light-colored wood, is a shortlived, small- to medium-sized tree (40 to 60 feet [12 to 18 meters] tall; 30 to 50 feet [9 to 15 meters] crown spread) (Fig. 1). Butternut's native range includes most of the northeastern United States and southern Canada from New Brunswick to Georgia, and west to Arkansas and Minnesota (Rink 1990; Dirr 1998) (Fig. 2). Butternut often grows in widely scattered clusters, with each cluster containing a few individual trees. It was never a highly abundant species (Schultz 2003), but for reasons that will be described later, it is even less common now than before. The former prevalence of--and appreciation for--butternut in the landscape is reflected evocatively by the many Butternut Hills, Butternut Creeks, and Butternut Lakes found across the eastern United States. Butternut is a member of the walnut family (Juglandaceae), which includes many familiar nut trees including eastern black walnut (Juglans nigra), Persian or English walnut (J. regia), pecan (Carya illinoinensis), and all the hickories (Carya spp.). How butternut relates to the other Figure 1. Researchers collect samples from a true butternut growing in walnuts remains a puzzle. Early taxonomy Daniel Boone National Forest, Kentucky. placed butternut in its own section within Juglans (Trachycaryon), but more recent treatone of the most winter-hardy, to USDA Zone 3 ments place it with Japanese walnut (J. ailanti(average annual minimum temperature -30 to folia) and Manchurian walnut (J. mandshurica) -40F [-34 to -40C]). in section Cardiocaryon (Manning 1978; FjellFood, Furniture, and Forage strom and Parfitt 1994), or with the New World Butternut has a long history of usefulness. walnuts (Rhysocaryon) (Aradhya et al. 2007). Native Americans extracted oil from the Butternut cannot hybridize with eastern black crushed nuts by boiling them in water, made walnut, but it can hybridize with Persian walnut syrup from the sap (Goodell 1984), and threw to form J. quadrangulata, and with Japanese butternut bark (which contains toxins) into walnut to form J. bixbyi (USDA-NRCS 2004). small streams to stun and capture fish. They Of all the walnuts, butternut is considered to be Butternut 3 BILL COOK, MICHIGAN STATE UNIvERSITy, BUGWOOD.ORG taught early European settlers how to make medicine from butternut bark, roots, and husks (Johnson 1884; Krochmal and Krochmal 1982). The inner bark of butternut and its nut hulls can be used to produce a yellow-brown dye. This dye was used most notably on some of the Confederate Army's Civil War uniforms, giving rise to the practice of referring to southern troops and their sympathizers as \"butternuts\" (Peattie 1950). Butternut is valued economically and ecologically today for its wood and edible nuts (Ostry and Pijut 2000) (Fig. 3). The sweet, oily, edible nuts are used in baked goods and are also popular for making maple-butternut candy. Butternuts were often planted near homes on farmsteads for the use of the nuts. There has been limited selection of butternuts for nut quality and production (McDaniel 1981; Goodell 1984; Miliken and Stefan 1989; Miliken et al. 1990; Ostry and Pijut 2000), but a few butternut cultivars with large nut size and superior ease of cracking (e.g., `Chamberlin' and `Craxezy') have been propagated, and some of these are available from commercial nurseries. The nuts are also an important food source for wildlife. In forests, butternut trees produce RINK, G. 1990 Figure 3. Butternut fruits have thick husks covered with sticky glandular hairs. Inside the husk is an edible nut enclosed in a thick, hard shell that is elaborated with eight prominent ridges (Brinkman 1974; Flora of North America Editorial Committee 1993+). NANCy ROSE Figure 4. Butternut wood samples: (clockwise from upper left) bark, slab-sawn, quarter-sawn, and cross-section (note darker brown heartwood). From the Ralph F. Perry wood collection at the Arnold Arboretum. Figure 2. The native range of butternut. seed at about 20 years of age, with good seed crops occurring every two to three years (Rink 1990). Open-grown trees, which benefit from more sun and less competition, can begin bearing as early as five years of age and bear annually under ideal conditions. The sapwood of butternut is light tan to nearly white and the heartwood is light brown (Fig. 4). The wood is moderately hard, but workable; it saws and carves easily, finishes well, and resembles black walnut when stained. The commercial availability of butternut wood is now extremely limited, but quality butternut 4 Arnoldia 66\/4 COURTESy OF GREy TOWERS NATIONAL HISTORIC SITE dead (Cummings-Carlson 1993; Cummings-Carlson and Guthmiller 1993). By the early 1990s butternut canker was reported in Canada (Davis et al. 1992), and butternut is now considered an endangered species in that country. In 1992, the state of Minnesota placed a moratorium on the harvest of healthy butternut on state lands, and butternut is considered a species of special concern in all United States National Forests. Although the origin of the fungus is uncertain (evidence suggests it may have come from Asia), it is believed to have been introduced into North America as a single isolate (Furnier et al. 1999). ButterFigure 5. The library at Grey Towers National Historic Site is paneled nut trees of all ages and sizes, regardless in butternut. of site conditions, can be infected. The wood commands a high market price today for spores of the fungus are spread by rain splash many uses including furniture, veneer, cabinets, and aerosols to adjacent trees where new infecpaneling, specialty products such as instrument tions originate at leaf scars, lateral buds, bark cases, interior woodwork, and fine woodworkwounds, and natural bark cracks. Perennial ing. The library of Grey Towers, a National cankers eventually develop on twigs, branches, Historic Site near Milford, Pennsylvania, and stems, and even the buttress roots (Tisserat and formerly the home of Gifford Pinchot, the first Kuntz 1983). Cankers can be seen most easily if chief of the United States Forest Service, is panthe bark is removed, revealing a sunken, ellipeled entirely with butternut (Fig. 5). tically-shaped region of dark brown to black stained wood, often with an inky black center A Deadly Disease Arrives and a whitish margin (Ostry et al. 1996) (Fig. 6). Sadly, a devastating canker disease has caused Cankers reduce the quality and marketability range-wide butternut mortality in recent of the wood, and the girdling effect of multiple decades and threatens the survival of the specoalescing cankers eventually kills a host tree. cies. Unusual stem cankers were first observed on butternuts in southwester n Wisconsin in 1967 (Renlund 1971). A pest alert announcing butternut decline was issued in 1976 (USDA 1976), and by 1979, the fungus responsible for butternut canker disease, Sirococcus clavigignenti-juglandacearum, was described as a new species (Nair et al. 1979). Surveys of butternut trees in Wisconsin in the 1990s revealed that 92% were diseased and 27% were Figure 6. Healthy butternut (left), and tree with bark removed showing cankers (right). PHOTO ON LEFT By KEITH WOESTE. PHOTO ON RIGHT USDA FOREST SERvICE FOREST HEALTH PROTECTION ST. PAUL ARCHIvE, USDA FOREST SERvICE, BUGWOOD.ORG Butternut 5 KEITH WOESTE While its spread to adjacent trees is understood, just how the fungus travels long distances to find new hosts remains a mystery. Several beetle species have been found on infected trees carrying fungal spores (Katovich and Ostry 1998; Halik and Bergdahl 2002), but it is not known which species (if any) carry spores over long distances. The fungus has also been found on the fruits of butternut and black walnut, causing lesions on the husks of both species (Innes 1998), which means that the movement of seeds can also spread the disease. Conservation and Restoration of Butternut There is no cure for butternuts once they become infected with butternut canker. In order to maintain butternut populations, conservationists must rely on a strategy of encouraging the growth of as many young, healthy trees as possible. The methods used include the management of regeneration (often by improving local habitats for seedling establishment) and reintroduction (for example, planting butternuts into suitable habitats from which they have been lost) (Ostry et al. 1994). Figure 7. Foresters identified this healthy butternut in a central Indiana forest. Butternut is a pioneer species, its seedlings require full sun to thrive (Rink 1990), percentage of the mature butternuts growand the presence of areas of exposed soil seems ing in the eastern forest are cankered, and to benefit its establishment (Woeste, personal infected trees have limited energy reserves observation). These factors explain why young to put towards flower and fruit production. butternuts tend to be found now on road-cuts, Because butternuts almost never self-pollinate steep terrain, fence-rows, old fields, clear-cuts, (Ross-Davis et al. 2008b), when a high percentwashouts, and the banks of swiftly flowing age of the trees in an area become diseased or streams. The management of most hardwood are killed, the number of potential mates can forests--both public and private--favors minbe reduced to the point that adverse genetic imal disturbance, so there are relatively few and demographic consequences become likely large, sunny openings for butternut seedlings to (Geburek and Konrad 2008). find a foothold. Browsing and antler rubbing by For all the above reasons and more, poor natdeer also limit the growth and survival of butural regeneration has been a hallmark of the ternut seedlings in the few sites sunny enough butternut canker epidemic (Ostry and Woeste to support regeneration (Woeste et al. 2009). 2004; Thompson et al. 2006). Until we learn Butternut canker, of course, also plays an how to effectively assist natural regeneration important role in reducing the natural regenof butternuts, reintroduction will be needed eration of butternut (Ostry et al. 1994). A high to restore butternut populations to the eastern 6 Arnoldia 66\/4 KEITH WOESTE population crash. Butternut collections must be conserved as living specimens growing in arboreta or other repositories because butternut seeds do not remain alive in long-term storage (even controlled-environment seed banks) unlike the seeds of many other species (Bonner 2008). Butternut can be propagated vegetatively by cuttings (Pijut and Moore 2002), through tissue culture (Pijut 1997; Pijut 1999), and by grafting. The ideal seed source for butternut reintroductions would be an orchard of genetically diverse, locally adapted, and canker-resistant butternut trees. Starting in the 1980s, a small group of scientists began identifying, grafting, and growing butter nuts that appeared healthy even though they were growing in locations with many dead or diseased trees (Ostry et al. 2003). It was assumed that these candidate trees had been exposed to the canker disease fungus, but because they remained healthy--or at least sufficiently healthy to continue to grow and reproduce--it was hoped that some of them would have genes for resistance to butternut canker. By the late 1990s, about 200 of these trees had been identified by Dr. Michael Figure 8. The trunk of a very old buart growing in central Indiana. Ostry of the USDA Forest Service Northforest. Reintroduction, whether by afforestaern Research Station in St. Paul, Minnesota, and tion (establishing plantations on old fields) or other colleagues. by supplemental planting in existing habitats, Butternut or Buart? requires a ready source of seeds. Seeds from By growing a large number of butternuts genetically diverse and locally adapted sources together in one location, Ostry and others were are preferred (Broadhurst et al. 2008). Because able to observe differences among these trees seed supplies from wild trees are so unreliable, that had not been obvious at the time of collecnumerous state and federal agencies as well tion. Differences in traits such as nut size and as private nurseries have worked over the past branch habit led him to wonder if some of the 20 years or so to document the location and collected butternuts were, in fact, buarts (Ostry health of butternut trees that could be used as and Moore 2008). A buart (pronounced bew-art), seed sources (Fig. 7). Others have collected and also called a buartnut, is the common name for grown butternut trees to provide seeds that will Juglans bixbyi (hybrids between butternut and be needed for reintroduction. the exotic Japanese walnut) (Fig. 8). Buarts were These collections constitute a germplasm well known among nut growing enthusiasts repository for butternut, a living bridge to the in the United States and Canada, but virtually future, and a method for preserving the genetic unknown by dendrologists and forest biologists diversity of the species in the face of a devastating Butternut 7 KEITH WOESTE (Ashworth 1969). Buarts had probably already been growing unnoticed in yards and orchards for a generation when they were first described by Willard Bixby in 1919 (Bixby 1919). Japanese walnuts were introduced into the United States around 1860 (Crane et al. 1937). In Japan, these walnuts were exploited as a food source by early tribal settlers (Koyama 1978), but never became an important commercial nut crop. By the late 1800s, Japanese walnuts had become popular among nut growers in the eastern United States because the kernels separate easily from the shell, and because some horticultural selections of Japanese walnut have an attractive and distinctively heart-shaped shell (Crane et al. 1937) (Fig. 9). Trees bearing heartshaped nuts became known as heartnuts (technically J. ailantifolia var. cordiformis), and the hybrid combination of butternut plus heartnut results in the common name \"buart\". Cultivars of heartnut have been selected and named (Ashworth 1969; Woeste 2004), but heartnuts never became a market success in the United States, perhaps because the nuts, while exotic in appearance, tend to be bland tasting. Although Japanese walnut never became popular as a nut crop, it gained a permanent foothold in the New World by intermating with butternut. Over time, as buarts became more common and as the gene pools of butternut and Japanese walnut intermixed, it became almost impossible and certainly impractical for most people to distinguish butternuts from buarts (Fig. 10). As early as 1919, Bixby (1919) found that \"[c]ertain Japan walnuts [are] so near like butternuts as to be readily mistaken for them. . . . [A]s far as the appearance of the nuts was concerned, the butternut could not be well separated from certain Japan walnuts.\" Buarts are remarkable hybrids. They stand out as exceptionally vigorous trees, sometimes exceeding 40 inches (102 centimeters) in diameter when mature (butternuts typically reach 12 to 24 inches [30 to 61 centimeters] in diameter). Buarts often bear enormous crops of nuts, and typically appear to be resistant to butternut canker (Orchard et al. 1982), although it is not certain that these trees truly are more resistant. It is easy to see why nut enthusiasts found buarts so attractive. Figure 9. A distinctively shaped heartnut (center) surrounded by nuts of other (non-heartnut) forms of Japanese walnut. All the nuts in the photo came from Japanese walnuts grown at the National Clonal Germplasm Repository for Juglans, in Davis, CA. PHOTO COURTESy SALLy WEEKS Figure 10. Butternut (top row) and hybrid nuts (bottom row) with the husks removed look nearly identical. As butternut populations dwindled and disappeared because of canker, buarts began to confound butternut conservation. Buarts were mistakenly identified as butternut survivors, and buarts planted in yards, parks, and cemeteries attracted seed collectors who gathered and sold the nuts to nurseries or through local markets, made them available through local conservation groups, or simply gave them away to friends and neighbors. Concerns about butternut's status in the forest caught some unaware because there were so many large, healthy \"butternuts\" (really buarts) growing in farmyards all over the countryside. It is likely that landowners have planted many more buarts than butternuts over the past 20 years, since so many of the 8 Arnoldia 66\/4 Figure 11. Summary of Characteristics Distinguishing Pure Butternut from Hybrid Butternuts. CHARACTERISTICS HABITAT BuTTERNuT Forests, occasionally as a grafted tree or wildling BuTTERNuT HyBRIDS Parks, forest edges, farmyards, urban areas, planted trees, orchards 1-yr-Twigs CuRRENT-yEAR STEm Olive green changing to red-brown near terminal, glossy, few hairs except immediately beneath terminal buds Beige in color; longer and narrower than hybrids, and the outer, fleshy scales more tightly compact. Vegetative buds are elongated (sometimes stalked) and somewhat angular, creamy white to beige in color Small, round, abundant, evenly distributed, sometimes elongating horizontally across the branch (perpendicular to the stem axis) Bright green to copper brown or tan, often densely covered with russet or tan hairs, especially near terminal buds. Pale green near terminal bud Pale green to tan or yellowish in color, wider and squatter than J. cinerea. Outer fleshy scales more divergent than butternut and often deciduous. Vegetative buds are rounded, and green to greenish brown in color. TERmINAl BuD lATERAl BuD lENTICElS large, often elongating laterally down the branch (parallel to the stem axis) on 1-yrwood, patchy distribution. On 3 and 4-yrwood, lenticels often form a diamond pattern as they become stretched both transversely and longitudinally Top edge almost always notched; often with large, exaggerated lobes Dark brown, medium brown or even light brown lEAF SCAR Top edge almost always straight or slightly convex; scar usually compact Dark brown PITH MaTure Tree BARK Varies from light grey and platy to dark grey and diamond patterned in mature trees. In older trees, fissures between bark ridges may be shallow or deep but are consistently dark grey in color. leaves yellow and brown by earlymid autumn, dehiscing in early to mid autumn. 512 cm in length at peak pollen shed One or two nuts per terminal in most clusters, sometimes 35, rarely more. Silvery or light grey, rarely darker. Fissures between bark ridges moderate to shallow in depth and often tan to pinkish-tan in color. lEAF SENESCENCE leaves often green until late autumn, dehiscing in late autumn or may freeze green on the tree. 1326 cm in length at peak pollen shed usually 3 to 5 per cluster, sometimes as many as 7. WOESTE ET AL. 2009 CATKINS NuT CluSTERS Butternut 9 remaining butternut trees have low vigor because of the effects of butternut canker and because butternuts, even when healthy, usually only produce a crop every two to three years (Rink 1990). For butternut, the existence of these hybrids presents something of a dilemma. On the one hand, buarts represent the dilution and potential loss of a distinctive native species with deep cultural connections and a complex quilt of ecological roles that evolved over many hundreds of thousands of years. On the other hand, hybridization is a common theme in plant evolution (Wissemann 2007), and for butternut, hybridization Figure 12. Twigs of butternut (top and bottom), Japanese walnut (upper middle,) could represent a way forward, espe- and buart (lower middle). The shape of the lenticels is characteristic of each type. cially if it is determined that all butternuts are completely susceptible to butternut Clonal Germplasm Repository in Davis, Calicanker (something that is far from certain at fornia, for comparison. this point). What role hybrids will play in butArmed with the best possible descriptions of ternut recovery remains to be seen. butternut and Japanese walnut, we had to conclude that trees with intermediate traits were Detailing the Differences buart hybrids. After examining a large number Whatever the possible uses of buarts, by 2003 of samples we developed a list of characters that it became clear to researchers that they needed can be used in combination to separate butterreliable mechanisms to distinguish buarts from nut and hybrids (Woeste et al. 2009) (Fig. 11). butternuts (McIlwrick et al. 2000; Ostry et al. After a few years of observing these traits in the 2003; Michler et al. 2005). The first task was field we have trained our eyes and now find that to describe the two parental species. Published most hybrids are fairly easy to spot, though for descriptions of the vegetative and reproductive more complicated cases a careful examination tissues of butternut, Japanese walnut, and the is needed to make a determination. (Fig. 12) hybrids are often brief, and based on an unknown At the same time, we began development of number of samples of unidentified provenance. a series of DNA-based tools for identifying butBy surveying published descriptions of butterternuts and hybrids (Ross-Davis et al. 2008a). nut, especially those made before the introducThe DNA markers are being used in both the tion of Japanese walnut to the United States or United States and Canada to identify true butbefore hybrids had an opportunity to become ternut seed sources. To understand the genetic widespread, a clearer picture of the morpholdiversity of butternut, we developed DNAogy of butternut and Japanese walnut emerged based markers called microsatellites, and used (Ross-Davis et al. 2008a). To verify our findthese to evaluate samples of butternuts from ings, we examined old butternut specimens at five locations spanning the upper south and the Herbarium of the Missouri Botanical Garmidwestern United States. To our relief, we den. These long-preserved samples provided learned that the genetic structure and neutral additional certainty that what we saw in the genetic diversity (diversity at the DNA level wild today matched what was collected over that is not associated with genes) of the current 100 years ago. We also obtained authenticated generation of large, standing butternuts was samples of Japanese walnut from the National quite similar to that of black walnut, a much KEITH WOESTE 10 Arnoldia 66\/4 KEITH WOESTE Figure 13. young butternut trees are screened for canker resistance at the Hardwood Tree Improvement and Regeneration Center in West lafayette, Indiana. more common related species (Ross-Davis et al. 2008b). This observation held out hope that it was not too late to begin to collect and preserve the genetic diversity of butternut. Armed with new DNA-based markers, and support from The Nature Conservancy and the USDA Forest Service State and Private Forestry, a small group of scientists and collaborators spent 2008 collecting butternut seeds as part of a long-term gene conservation program. A permanent home for the seedlings that will grow from these seeds is envisioned in western Iowa, sufficiently distant from sources of butternut canker it is hoped, to ensure the collection will be safe. These trees represent one of several collections that will reconstitute the future for butternut. A final note of good news is that an evaluation of candidate canker-resistant butternuts using our DNA-based methods confirms that many of the trees are truly butternuts and not hybrids (Woeste, unpublished data). Recently, pathologists proposed protocols for inoculating and testing candidate trees to determine if these are truly resistant to butternut canker (Ostry and Moore 2008) (Fig. 13). If future pathology studies demonstrate that some candidate trees contain useful levels of resistance to butternut canker, an aggressive program of breeding will be undertaken to transfer the resistance genes into butternuts from all across the species' range. The goal will be to produce seed orchards of genetically diverse, regionally adapted, diseaseresistant butternuts for reintroduction to areas of the eastern forest where butternut has disappeared. Learning how to reintroduce and sustain viable populations of trees into habitats from which they have been lost remains an important and ongoing challenge (Broadhurst et al. 2008; Geburek and Konrad 2008). Butternut 11 References Aradhya, M., D. Potter, F. Gao, and C. Simon. 2007. Molecular phylogeny of Juglans (Juglandaceae): a biogeographic perspective. Tree Genetics and Genomes 3: 363378. Ashworth, F.L. 1969. Butternuts, Siebold (Japanese) Walnuts, and Their Hybrids. pp. 224231, In: Handbook of North American Nut Trees. 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Cummings-Carlson, J. and M. Guthmiller. 1993. Incidence and severity of butternut canker in Wisconsin in 1976 and 1992. Phytopathology 83: 1352. Davis, C.N., D.T. Myren, and E.J. Czerwinski. 1992. First report of butternut canker in Ontario. Plant Disease 76: 972. Dirr, M.A. 1998. Manual of Woody Landscape Plants: Their Identification, Ornamental Characteristics, Culture, Propagation and Uses. 5th ed., Stipes Publishing, Champaign. Fjellstrom, R.G. and D.E. Parfitt. 1994. Walnut (Juglans spp.) genetic diversity determined by restriction fragment length polymorphisms. Genome 37: 690700. Flora of North America Editorial Committee. 1993+. Flora of North America North of Mexico, 7+ vols. New york and Oxford. Furnier, G.R., A.M. Stolz, R.M. Mustaphi, and M.E. Ostry. 1999. Genetic evidence that butternut canker was recently introduced into North America. Canadian Journal of Botany 77 (6): 783785. Geburek, T. and J. Konrad. 2008. Why the conservation of forest genetic resources has not worked. Conservation Biology 22 (2): 267274. Goodell, E. 1984. Walnuts for the northeast. Arnoldia 44 (1): 319. Halik, S. and D.R. Bergdahl. 2002. Potential beetle vectors of Sirococcus clavigignenti-juglandacearum on butternut. Plant Disease 86: 521527. Innes, L. 1998. Sirococcus clavigignenti-juglandacearum on butternut and black walnut fruit. pp. 129 132, In: Foliage, Shoot, and Stem Diseases of Trees; Proceedings of the International Union of Forest Research Organizations. LaFlamme, G.; Berube, J.A.; Hamelin, R.C. (eds.) Working Party 7.02.02; 1997 May 2531; Quebec City, Canada. Johnson, L. 1884. Manual of the Medical Botany of North America. W. Wood and Co., New york. Katovich, S.A. and M.E. Ostry. 1998. Insects associated with butternut and butternut canker in Minnesota and Wisconsin. The Great Lakes Entomologist 31: 97108. Koyama, S. 1978. Jomon subsistence and population. Senri Ethnological Studies 2: 1246. Krochmal, A. and C. Krochmal. 1982. Uncultivated nuts of the United States. USDA Forest Service Agriculture Information Bulletin 450. Manning, W.E. 1978. The classification within the Juglandaceae. Annals of the Missouri Botanical Garden 65: 10581087. McDaniel, J.C. 1981. Other walnuts including butternut, heartnut, and hybrids. pp. 98110, In: Nut Tree Culture in North America. R.A. Jaynes (ed.), Northern Nut Growers Association, Hamden. McIlwrick, K., S. Wetzel, T. Beardmore, and K. Forbes. 2000. Ex situ conservation of American chestnut (Castanea dentata [Marsh.]) and butternut (Juglans cinerea L.), a review. Forestry Chronicle 76: 765774. Michler, C. H., K.E. Woeste, P.M. Pijut, D.F. Jacobs, R. Meilan, and M. Ostry. 2005. Improving disease resistance of butternut (Juglans cinerea) a threatened fine hardwood: a case for single tree selection through genetic improvement. Tree Physiology 26: 113120. Milikan, D.F. and S.J. Stefan. 1989. Current status of the butternut, Juglans cinerea L. Annual Report of the Northern Nut Growers Association 80: 5254. Milikan, D.F., S.J. Stefan, and K.S. Rigert. 1990. Selection and preservation of butternut, Juglans cinerea L. Annual Report of the Northern Nut Growers Association 81: 2225. Nair, v.M.G., C.J. Kostichka, and J.E. Kuntz. 1979. Sirococcus clavigignenti-juglandacearum: an undescribed species causing canker on butternut. Mycologia 71: 641646. 12 Arnoldia 66\/4 Orchard, L.P., J.E. Kuntz, and K.J. Kessler, Jr. 1982. Reaction of Juglans species to butternut canker and implications for disease resistance. pp. 2731, In: Proceedings of Conference on Black Walnut for the Future. General Technical Report NC-74. USDA Forest Service, North Central Forest Experiment Station, St. Paul, MN. Ostry, M.E., M.E. Mielke, and D.D. Skilling. 1994. Butternut--Strategies for managing a threatened tree. General Technical Report NC-165. USDA Forest Service, North Central Forest Experiment Station, St. Paul, MN Ostry, M.E, M.E. Mielke, and R.L. Anderson. 1996. How to identify butternut canker and manage butternut trees. HT-70. USDA Forest Service, Northeastern Area State and Private Forestry. Ostry, M.E. and P.M. Pijut. 2000. Butternut: An underused resource in North America. HortTechnology 10 (2): 302306. Ostry, M.E., B. Ellingson, D. Seekins, and W. Ruckheim. 2003. The need for silvicultural practices and collection of butternut germplasm for species conservation. pp. 551555, In: Proceedings of 13th Central Hardwood Forest Conference. General Technical Report NC-234. USDA Forest Service, North Central Research Station, St. Paul, MN Ostry, M. E. and K. Woeste. 2004. Spread of butternut canker in North America, host range, evidence of resistance within butternut populations and conservation genetics. pp. 114120, In: Black Walnut in a New Century, Proceedings of 6th Walnut Council Research Symposium. Michler, C.H. et al., eds.; 2004 July 2528, Lafayette, IN. General Technical Report NC-243. USDA Forest Service, North Central Forest Experiment Station, St. Paul, MN. Ostry, M.E. and M. Moore. 2008. Response of butternut selections to inoculation with Sirococcus clavigignenti-juglandacearum. Plant Disease 92: 13361338. Peattie, D.C. 1950. A Natural History of Trees of Eastern and Central North America. Houghton Mifflin, Boston. Pijut, P.M. 1997. Micropropagation of Juglans cinerea L. (Butternut). pp. 345357, In: Biotechnology in Agriculture and Forestry, vol.39, High-Tech and Micropropagation, Section III.4:, Bajaj, y.P.S. (ed.). Springer-verlag, Berlin, Heidelberg, New york. Pijut, P.M. 1999. Somatic embryogenesis from immature fruit of Juglans cinerea. pp. 415429, In: Somatic Embryogenesis in Woody Plants, vol.4, Section B: Jain. S.M., Gupta, P.K., and Newton, R.J. (eds.)., Kluwer Academic Publishers, The Netherlands. Pijut, P.M. and M.J. Moore. 2002. Early season softwood cuttings effective for vegetative propagation of Juglans cinerea. HortScience 37 (4): 697700. Renlund, D.W. 1971. Forest pest conditions in Wisconsin. In: Annual Report of the Wisconsin Department of Natural Resources, Madison, WI. Rink, G. 1990. Juglans cinerea L., Butternut. pp. 386390, In: Silvics of North America, vol 2. Hardwoods. R.M. Burns and B.H. Honkala (Tech. Coords.). USDA Forest Service Agriculture Handbook 654, Washington, DC. Ross-Davis, A., Z. Huang, J.R. McKenna, M.E. Ostry, and K. Woeste. 2008a. Morphological and molecular methods to identify butternut (Juglans cinerea) and butternut hybrids: relevance to butternut conservation. Tree Physiology 28: 11271133. Ross-Davis, A., M.E. Ostry, and K. Woeste. 2008b. Genetic diversity of butternut (Juglans cinerea) and implications for conservation. Canadian Journal of Forest Research 38 (4): 899907. Schultz, J. 2003. Conservation assessment for butternut or white walnut (Juglans cinerea L.). USDA Forest Service, Eastern Region. On-line at www.fs.fed. us\/r9\/wildlife\/tes\/ca-overview\/docs\/plant_ juglans_cinera-Butternut2003.pdf Thompson, L.M., F.T. van Manen, S.E. Schlarbaum, and M. DePoy. 2006. A spatial modeling approach to identify potential butternut restoration sites in Mammoth Cave National Park. Restoration Ecology 14: 289296. Tisserat, N. and J.E. Kuntz. 1983. Dispersal gradients of conidia of the butternut canker fungus in a forest during rain. Canadian Journal of Forest Research 13 (6): 11391144. USDA. 1976. Butternut decline. Pest Alert. USDA Forest Service, Misc. Publication Newtown Square, PA, Northern Area State and Private Forestry. USDA-NRCS. 2004. The PLANTS Database, version 3.5 (http:\/\/plants.usda.gov). National Plant Data Center, Baton Rouge. Wissemann, v. 2007. Plant evolution by means of hybridization. Systematics and Biodiversity 5: 243253. Woeste, K. 2004. An On-line Database of Juglans Cultivar Names and Origins. HortScience 39: 1771. Woeste, K., L. Farlee, M. Ostry, J. McKenna, and S. Weeks. 2009. A forest manager's guide to butternut. Northern Journal of Applied Forestry. Keith Woeste and Paula M. Pijut conduct research at the USDA Forest Service Northern Research Station Hardwood Tree Improvement and Regeneration Center in West Lafayette, Indiana. "},{"has_event_date":0,"type":"arnoldia","title":"A Closer Look at Fungi in the Arnold Arboretum","article_sequence":2,"start_page":13,"end_page":21,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25455","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14eb76d.jpg","volume":66,"issue_number":4,"year":2009,"series":null,"season":null,"authors":"Richardson, Kathryn","article_content":"A Closer Look at Fungi in the Arnold Arboretum Kathryn Richardson W alk into the Arnold Arboretum and you'll see a beautiful and historically important collection of trees and shrubs. With a closer look, other organisms become visible, including fungi. The world of fungi is vast, and many members of this diverse kingdom are found throughout the Arboretum. In an informal survey from spring through fall of 2008, Arboretum staff reported over 100 fungal sightings, and positive identifications were made for 40 species. A dozen species noted in the survey are profiled starting on page 15. Fungi ID PrecIsely identifying fungi can be challenging even for experts. Many morphological features from growth habit to the size and color of spores provide clues for identification. Often minute details are needed to confirm species identity. correct identification is critical when considering fungi as food, since the fungi kingdom contains species that are deliciously edible and others that are deadly poisonous. Though often stated, it's worth repeating: Never consume any mushroom without being absolutely sure of its identity. susAN HArdy BrOWN Plants + Fungi The association between plants and fungi is sometimes beneficial, sometimes harmful, and sometimes a bit of both. Fungi can be indicators of a tree's declining health, but in other cases--such as mycorrhizae which aid roots with nutrient uptake--they are physiologically beneficial to plants. Identifying fungi and understanding their physiology is an important aspect of interpreting the health of the Arboretum's collections. The Arnold Arboretum hosts an unknown number of fungi, many of which have specific relationships with certain plants. When a fungus is found growing on an accessioned tree, horticultural staff attempt to make an accurate identification, and often those known relationships aid in the identification process. For example, a shiny bracket fungus growing on eastern hemlock (Tsuga canadensis) is probably the hemlock varnish shelf (Ganoderma tsugae), and an oak tree with a massive cluster of sulphur-yellow mushrooms growing from its roots is likely to be chicken of the woods (Laetiporus sulphureus). These fungi may live in their hosts for several years. As they feed, these fungi cause wood decay and often weaken the The distinctive lamellae (gills) of this mushroom mark it as a member of the phylum Basidiomycota, but much more information is needed to determine its exact species. 14 Arnoldia 66\/4 structural integrity of the tree. damage from fungi also weakens the tree's defenses and may enable entry for other pests and diseases which cause further injury. Arboretum trees flagged with potentially fatal fungi are carefully observed and notes are kept within the Arboretum's collections database. If the tree begins to decline, this information is useful in making a decision on its treatment or removal. ing for miles), or within wood or other hosts. Mycelia grow even when no fruiting bodies are present, so the extent of damage caused by a fungus in the tree before the emergence of the fruiting body is sometimes unclear. The Fungal Life Fungi are neither plant nor animal and are placed in their own kingdom, though historically this was not always the case. Taxonomists initially placed fungi in the plant kingdom (Plantae) because, plants and fungi are both sessile (not free-moving) and have cell walls. However, fungi lack chlorophyll (and thus cannot make their own food via photosynthesis) and have walls made of chitin, not cellulose as seen in plants. Fungi are closely related to animals and bacteria and were once placed in the animal kingdom (Animalae), but fungi are not motile. Fungi proved to be unique life forms deserving their own kingdom. Fungi cannot produce their own food and thus must acquire the nutrients they need from their hosts or substrates. Based upon their nutritional needs, fungi fall into three categories: saprobes, parasites, and mutualists. saprobic fungi feed on dead organic materials and serve as the scavengers of the kingdom by recycling carbon, nitrogen, and other essential elements back into the soil. Parasitic fungi feed on living organisms and often harm them in some way. \"Parasitize\" sounds threatening, but the reality is that fungal parasites do not typically destroy their host quickly and may be present for some time before the host shows decline. Mutualistic fungi have a beneficial relationship with other living organisms. examples of mutualists include lichens (fungi plus algae or cyanobacteria) and mychorrhizae (fungi and plant roots). Around 90% of all living trees have a mycorrhizal relationship with fungi. The presence of fungal fruiting bodies on trees indicates that the fungus has reached the spore production stage of its life cycle. When released spores land on a substrate and germinate, threadlike hyphae grow and combine to form mycelia, the vegetative growth of fungi. Mycelia grow underground (sometimes spread- Fungi to Know The largest groups of fungi are found in the phylum Basiodiomycota, often inclusively called the basidiomycetes. Many familiar fungi such as cap-and-stipe (stalk) mushrooms (including the cultivated \"white button mushroom\" [Agaricus bisporus] found in grocery stores), brackets, and puffballs are placed in this group. It is safe to say that if you are looking at a fungus that has either rib-like gills (lamella) or tiny pores on the underside of the cap, it's a basidiomycete. The microscopic rust and smut fungi are also basidiomycetes. Basidiomycota produce basidiospores which have a single haploid nucleus. When these spores germinate they produce long, branching hyphae with a single nucleus in each compartment (area between cell walls). When two compatible hyphal strands come into contact with each other they unite to form a hyphal strand that now houses two nuclei in each compartment. A basidiomycete will spend most of its life in the vegetative mycelial stage until environmental cues, such as rain or temperature change, cause the growth of fruiting bodies (basidiocarps). Many basidiomycetes are decomposers, but others have a mycorrhizal partnership with forest trees. Another fungal phylum, Ascomycota, includes the sac fungi or spore shooters. unlike basidiomycetes that have structures (basidiophores) that drop spores from their fruiting bodies, ascomycetes have spores in sacs located within a structure called an ascocarp or ascomata. The spores are \"shot\" out of their sacs and dispersed into the air. sac fungi are also decomposers and recyclers of organic matter. Many ascomycetes are parasitic including those that cause dutch elm disease and chestnut blight. Ascomycetes include yeasts, which are used to make beer and wine, as well as mycelial fungi such as morels and black knot. Another interesting example of an ascomycete is the fungus that causes ergot, a damaging disease of grain crops. ergot fungus contains a compound Fungi 15 that is a precursor to the hallucinogen lsd. Though not proven, it has been suggested that ergot poisoning was a potential cause of the hysteria that led to the salem witchcraft trials in the late 1600s. Although no longer classified in the fungi kingdom, Myxomycetes (slime molds) are also mentioned here since they resemble fungi, are common at the Arboretum, and elicit many questions from visitors (see page 21). A Sampler of Arboretum Fungi Here are a dozen interesting fungi--plus one slime mold--that were found in last year's informal survey of fungi at the Arnold Arboretum cOurTesy OF PAulA desANTO Pheasant's-back Polypore or Dryad's Saddle (Polyporus squamosus) In the spring of 2008, a very large pheasant's-back polypore appeared on a venerable cucumbertree magnolia (Magnolia acuminata, accession 15154-e) near the main entrance of the Arnold Arboretum. This magnolia has survived for over 100 years, but time has taken its toll and the doors are now open for various organisms, including fungi, to invade. Growing out of an old limb-removal wound on the magnolia's trunk was an impressive bracket with a uniquely patterned cap. This species, the pheasant's-back polypore, can grow to 24 inches (61 centimeters) in diameter. It appears growing on stumps and dead hardwood trees in spring in the northeastern united states. It is easily recognized by its fan-shaped, tan to creamy yellowish cap with an array of brown scales that look like pheasant feathers-- thus the species' common name. (Another common name, dryad's saddle, refers to its potential use by the tree-dwelling nymphs known as dryads in Greek mythology.) The white underside of the cap is dotted with thousands of small pores (polyporus means \"many pores\"). It was sad to see this particular polypore because it indicates that this magnolia's life is coming to an end. The Arboretum's horticultural staff had noted the tree's decline before the emergence of this polypore, but its presence told us more about the health of this tree. The pheasant's-back polypore is typically saprobic on dead trees but it can also parasitize the heartwood of living trees such as this magnolia. This polypore fungus had been living in this tree for an unknown period of time before it produced this fruiting body; the extent of internal rot is uncertain but the tree will continue to be monitored closely. 16 Arnoldia 66\/4 HAruTA OVIdIu, uNIVersITy OF OrAdeA, BuGWOOd.OrG Witch's Butter (Tremella mesenterica) Witch's butter is a member of the phylum Basidiomycota, but does not have the traditional cap and stem as do some other fungi in this group. This fungus is yellow to orange in color and appears as wavy, gelatinous folds. It can dry out to the point of appearing dead, but will rehydrate readily with rainfall or other applied water. Another interesting fact about witch's butter is that it feeds on other fungi, not on wood. It is often seen growing on downed logs or dead branches, where it parasitizes wood-decaying fungi. Witch's butter is widely distributed in temperate regions in North America, europe, Asia, and Australia. The name \"witch's butter\" comes from several european legends. One states that if the fungus was found growing near a home's entrance or front gate, then the homeowner had been hexed by a witch. The spell could be broken by plunging a pin into the fungus, causing the witch to feel the pinpricks, which in turn would cause her to return to remove the spell and the fungus. A legend of swedish origin blames this fungus on a witch's cat. The cat, sent out to steal food from the neighbors, would gorge itself and then vomit \"witch's butter\" on the gardens, fences, gates, and homes of unsuspecting people. The name \"witch's butter\" is sometimes applied to any of a number of jelly-like fungi. NIMA sAMIMI The Stinky Squid (Pseudocolus fusiformis) The stinky squid is a basidiomycete belonging to the Phallaceae, a family of fungi commonly known as stinkhorns. The stinky squid certainly lives up to its name both in scent and appearance--in late August 2008 this stinkhorn created quite a horrible smell in the Arnold Arboretum when it appeared in a few beds in the leventritt shrub and Vine Garden as well as in densely planted areas on Peters Hill. common to eastern North America, this species of stinkhorn has a fantastic appearance. Beginning its reproductive life as an egglike structure with white rhizomorphs attached to the base, its fruiting body quickly emerges, displaying three to five tapering arms. The arms may be free-standing or fused together at the tips, and are yellow towards the base and reddish orange towards the apex. It stands 1 to 3 inches (3 to 7 centimeters) in height with dark green spores lining the inner sides of its arms. (A broken-off fruiting structure is seen here.) There's no question about how this fungus received its common name: it looks like a squid and has the odor of rotting flesh. stinkhorns, including stinky squid, Fungi 17 disperse their spores by attracting flies and other insects which land on the fungus and feed on the stinky slime. In the process, the insects collect spores on their bodies as well as ingesting them, then spread the spores to new locations. Common Oyster Mushroom (Pleurotus ostreatus) The common oyster mushroom--a familiar edible mushroom that can be found in grocery stores--is common in the Arboretum and appeared in large numbers last spring. Oyster mushroom species are typically found in the fall, winter, and early spring, though they are also sometimes seen in the summer under the right conditions. They grow on dead hardwoods and, less often, on conifers, and also on some living trees. Oyster mushrooms grow in dense clusters, have light brown to off-white caps, and display prominent, elongated white gills. An interesting fact about species in this mushroom genus is that they are carnivorous; they trap, kill, and eat living organisms such as nematodes and bacteria in addition to the more typical fungus function of decomposing wood. JOsePH O'BrIeN, usdA FOresT serVIce, BuGWOOd.OrG Chicken of the Woods (Laetiporus sulphureus) chicken of the woods belongs to the genus Laetiporus, which fairly recently has been separated into several species based on dNA analysis. When I first began identifying chicken of the woods in the Arboretum I assumed it was Laetiporus sulphureus, but most turned out to be the very similar-looking species Laetiporus cincinnatus. Both species have the common name \"chicken of the woods\" and are popular edibles for mushroom hunters. They are readily identified because of their bright yellow to orange color and appearance as masses or rosettes of wavy, blunt-rimmed plates. They are widely distributed east of the rocky Mountains and often grow as parasites or saprobes on oaks (Quercus spp.). The main difference between the two species is the location of their fruiting bodies; Laetiporus sulphureus usually grows on tree stumps while L. cincinnatus grows from the roots of the infected host, giving the appearance that it is growing terrestrially. The common name is appropriate for several reasons. The flesh of the caps is yellowish in color, almost like raw chicken. Also, the taste and texture of this fungus, when cooked, reportedly are similar to cooked chicken. NANcy rOse 18 Arnoldia 66\/4 rOBerT MAyer NANcy rOse Turkey Tail Fungus (Trametes versicolor) The turkey tail fungus is one of the most commonly seen bracket fungi, occurring on dead trees in temperate zone forests all over the world. Turkey tail fungus is saprobic on dead hardwoods and can sometimes completely cover trunks and branches. A decomposer of wood, this fungus will sometimes work away for hundreds of years on a single host. Turkey tail fungus is a polypore, having pores instead of gills, and has a hard exterior instead of the fleshy ones seen in traditional mushrooms. It is aptly named, displaying concentric colored bands that resemble a fanned turkey's tail. The colors of turkey tail fungus can vary, but the bands commonly appear in shades of white, brown, and tan, sometimes with more colorful bands in orange, cinnamon, or bluish tones. A close look reveals dense, downy hairs on the bracket's upper surface. usdA FOresT serVIce-NOrTH ceNTrAl reseArcH sTATION ArcHIVe, BuGWOOd.OrG Shaggy Mane Mushroom (Coprinus comatus) The shaggy mane--a type of inky cap mushroom--is readily observed from mid-spring to late summer. Found on lawns, in mulched beds, and in forests, the shaggy mane performs as one of nature's recyclers, feeding on soil, forest litter, decaying wood, and even dung. It slowly decomposes the organic matter on which it feeds. One characteristic that makes this mushroom interesting is its method of spore dispersal. When the spores begin to mature, the shaggy oval cap begins to curl, becoming bellshaped, as the gills deliquesce (liquefy). This gives the spores maximum exposure to the wind, which then transports the spores to new locations. The gills will continue to liquefy until they are virtually gone, leaving a flat, almost transparent cap. True to the name, the liquefied gills of this and other inky caps can be used as a semi-permanent ink. Bark Mycena (Mycena spp.) There are many tiny, often-overlooked mushrooms growing in the Arboretum including several in the genus Mycena. This genus contains hundreds of species distributed worldwide. Most Mycena species are very small and have bell-shaped caps on slender stipes. Walking along Meadow road I came across an old painted maple (Acer mono) covered with these tiny mushrooms. Gray-brown in color with caps no larger than a few millimeters in diameter they covered the bark of this Fungi 19 maple along with moss and lichens. At first it seemed sad to see such a fantastic old tree covered with mushrooms, but these fungi do not harm the tree. Bark Mycena live on the outer layer of a tree, feeding on the dead bark. They never move to the living layers of the tree and thus do no harm. Bird's-Nest Fungi (species in several genera including Crucibulum and Cyathus) Bird's-nest fungi are a group of unusual fungi in the order Nidulariales (\"nidula\" means small nest). They are very common in the Arboretum and can be found growing in almost every mulched bed as well as on debris in natural woodland areas. These harmless fungi are saprobic on substrates such as dead wood (including woodchips), leaves, and dung. They often grow in large expanses. The common name describes these fungi perfectly. The mature peridia (fruiting bodies) resemble tiny nests. These nests contain tiny egglike peridioles which contain spores. several species of bird's-nest fungi grow at the Arboretum and can be differentiated by the color, size, shape, and texture of their peridia as well as by the color of their peridioles which can vary from white to black with several shades of gray and brown in between. Bird's-nest fungi exhibit an interesting spore dispersal method. When it rains, water droplets splash the \"eggs\" (peridioles) out of the nest and into the air. When this happens, a cord which attaches the egg to the nest breaks free and elongates. When the egg lands on nearby substrates the cord sticks and secures the egg to its new site. rOBerT MAyer Hen of the Woods (Grifola frondosa) Hen of the woods is a popular edible mushroom with sweet-tasting flesh. The clustered caps of this fungus resemble the ruffled feathers of a hen, and a full-grown specimen can reach a foot or more in diameter and weigh as much as 40 pounds (18 kilograms). Hen of the woods is commonly found growing on oak (Quercus spp.) trees from either the trunk or roots. This mushroom is a parasite and will cause damage over time. It causes white rot which can compromises the structural integrity of the roots. A weakened root system can prove disasterous for a tree in wind storms, since lack of solid anchorage may allow the tree to topple over. NANcy rOse 20 Arnoldia 66\/4 NANcy rOse Black Knot Fungus (Apiosporina morbosa) Black knot fungus is visible on several cherry trees by the Arboretum's Forest Hills gate. Black knot can infect a number of cherry and plum species (Prunus spp.). This ascomycete is a harmful fungus that damages both the health and appearance of its host. The visible part of this fungus, a black gall, is the result of the fungus disrupting the normal growth of the twig. Galls form at the site of infection. Black knot galls look something like burned marshmallows on a stick and may eventually grow to a foot in length if left unchecked. Inside the galls are perithecia which produce ascospores, which, after overwintering in the gall, are ejected in the spring when warmer temperatures and adequate moisture arrive. The ascospores are then carried by wind and water to new host sites. Infection occurs on new plant growth and wounded tissues. These ascospores are able to penetrate through the green tissue of new growth and quickly begin to grow. New galls are brown, and can easily go unnoticed until the following year when they continue to grow and turn black. The galls continue to grow every year and the infection continues to spread further down the branch. Older galls often harbor borers which can cause even more problems for infected trees. All trees at the Arnold Arboretum with black knot galls present are monitored. When a gall is found the infected branch is removed while the fungus is still dormant. This slows further spread on the host tree and also reduces the spread of infection to other trees. rOBerT MAyer Hemlock Varnish Shelf (Ganoderma tsugae) The Arboretum's Hemlock Hill offers visitors a chance to see the interaction between a fungus and a specific type of tree. The hemlock varnish shelf (Ganoderma tsugae) has a preference for conifers and specifically for hemlocks (Tsuga spp.). It is found on living and fallen trees on Hemlock Hill and was also reported growing in a mulch bed along Meadow road. If seen growing on a living hemlock it is safe to say that the tree is not in perfect health. Fungi 21 The hemlock varnish shelf is a beautiful polypore. Its hard, shiny cap is dark red to reddish brown, sometimes with prominent concentric zones. young specimens may show white and yellow segments also. This annual mushroom grows individually or, less commonly, in limited clusters. This species is closely related to the more common taxa Ganoderma lucidum, (sometimes known as reishi or lingzhi); extracts of both have been used in herbal medicine. susAN HArdy BrOWN Dog Vomit Slime Mold (Fuligo septica) I have had Arboretum visitors ask me about \"the lumpy yellow (or tan) stuff in the mulch bed that looks like vomit.\" Well, that's the descriptively named dog vomit slime mold, commonly seen in planting beds mulched with wood chips. Fuligo septica is a type of Myxomycetes, so not a true fungus. It is a plasmodial slime mold; this means that the \"vomit\" is actually a huge single cell containing millions of nuclei. dog vomit slime mold is motile, but moves quite slowly. It is not harmful to animals or plants and usually vanishes in a short period of time. This species and similar slime molds feed on bacteria, fungal spores, and smaller protozoa found on wood chips. slime molds feed much like an amoeba feeds; they ingest their food and then digest it (unlike fungi, which digest and then ingest). If conditions are favorable, these slime molds will produce reproductive structures (sporangia) that produce spores. When conditions are unfavorable (loss of food, dry conditions), the plasmodium will form hard, dormant, protective structures called sclerotia. Inside the sclerotia the plasmodium will divide into \"cells\" containing up to four nuclei. When conditions become favorable each \"cell\" will form a new plasmodium. dog vomit slime mold is primarily an aesthetic problem in mulched garden beds. It can be physically removed, but more is likely to return. so, before panicking and taking your dog to the veterinarian, take a closer look and consider that that stuff is likely just Fuligo septica working away at cleaning the mulch. Acknowledgments Thanks to susan Hardy Brown, Nima samimi, eric youngerman, Bob ervin, Marc devokaitis, Nancy sableski, and all staff for their help in surveying and photographing fungi at the Arboretum, and to don Pfister for reviewing this article. Kathryn richardson is a curatorial Assistant at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"American Chestnuts in the 21st Century","article_sequence":3,"start_page":22,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25456","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14ebb26.jpg","volume":66,"issue_number":4,"year":2009,"series":null,"season":null,"authors":"Anagnostakis, Sandra L.","article_content":"American Chestnuts in the 21st Century Sandra L. Anagnostakis N ew England was heavily forested in 1600, and American chestnut (Castanea dentata) was commonly found in Connecticut and Massachusetts woodlands (Cogbill et al. 2002). At that time, American chestnut was abundant throughout its native range from southern Maine to northern Georgia, all along the Appalachian Mountains (Saucier 1973). In the following centuries, European settlers cleared land for farming and cut trees for fuel, and the forest cover was greatly reduced by 1850. This was followed by the introduction of coal as a fuel, which was brought easily to New England by the railroads. Once wood was no longer being harvested for fuel, and more fields were left fallow as people abandoned farms and moved west or into the cities, the trees started to take back their habitats. When hardwood forests were harvested and left to resprout, the chestnuts grew faster than the oaks and maples with which they shared the land, and the number of chestnut trees greatly increased. Many woodlots became nearly pure stands of chestnut. A bulletin issued by the Connecticut Experiment Station in 1906 stated that regenerating hardwood forests covered most of the wooded area of Connecticut and \"the most important tree of this type is the chestnut which constitutes fully one-half of the timber\" (Hawes 1906). Forest surveys done at the turn of the last century show that there were about 130 million mature American chestnut trees in Connecticut alone. These stands of chestnut trees were valued because chestnut is a strong wood that resists rotting. Chestnut was used extensively for framing and woodwork, and was also essentially the only wood used for telephone poles and most of the railroad ties laid as rail lines pushed westward (Pierson 1913). The Blight Arrives The fungal pathogen causing chestnut blight disease (now called Cryphonectria parasitica) was introduced into the United States in the late 1800s on Japanese chestnut trees. The disease was spread up and down the east coast by mail-order sales of infected trees (Anagnostakis 2001, http:\/\/www.ct.gov\/caes\/cwp\/view. asp?a=2815&q=376754). In 1908 chestnut blight disease started killing American chestnut trees in Connecticut (Clinton 1912), and SAUCIEr 1973 Native range of American chestnut (Castanea dentata) in Eastern North America. American Chestnut 23 FrOM USDA FOrEST SErvICE BUllETIN 96 (1912) A pure stand of American chestnut in Connecticut in 1910. infections were reported in Cape Cod, Wellesley, and Pittsfield, Massachusetts (Metcalf and Collins 1909). Chestnut blight disease has reduced American chestnuts to understory shrubs, which die back, sprout from the base, die back, and sprout again. This fungus is now present throughout the original range of C. dentata, and has spread to many of the Midwestern locations where chestnuts were planted. Chestnut Breeding Chestnut trees are monoecious and bear separate male and female flowers on the same tree. As with many fruit trees, they must be crosspollinated for fully formed nuts to develop. Without cross-pollination, burs with small, flat nuts comprised of all-female tissue are all that form. Although the size of the nuts formed is completely dependent on the female parent, the pollen parent influences the flavor of the nuts (Anagnostakis 1995a, Anagnostakis and Devin 1998). Growers interested in getting nuts as large as those of Japanese or European chestnut but with the superior flavor of American chestnuts started creating hybrids in the late 1800s. After chestnut blight disease began killing timber 24 Arnoldia 66\/4 DisEAsE NoT REpoRTED DisEAsE NoT BAD DisEAsE BAD This map shows the presence of chestnut blight disease in Connecticut in 1908. SANDrA ANAGNOSTAkIS Blight canker on an American chestnut tree; note the dead, sunken bark and lumps of fungal tissue that have broken through the surface where they will form spores. chestnut trees--and Asian chestnut trees were seen to be resistant to the disease-- it was hoped that new hybrids could be developed that combined the upright, timber-producing form of American chestnut with the Asian species' resistance to blight. Arthur Graves, a plant pathologist in Connecticut, began crossing blightresistant Asian trees and susceptible American trees in 1930. He then tested these hybrids for resistance to chestnut blight disease (Graves 1937). He was soon joined by Donald Jones of the Connecticut Agricultural Experiment Station (CAES), who was a renowned geneticist with a great interest in chestnut. Many of those original hybrids are still alive, and CAES now has what is probably the finest collection of species and hybrids of chestnut in the world. These were planted on land left to the State of Connecticut by Graves, and at the CAES farm, both located in Hamden, Connecticut. Trees with two forms are being chosen from our continuing breeding efforts at CAES: tall, straight trees with limited energy put into forming nuts but very well-suited for timber production, and short, spreading trees with maximum energy put into forming large, good-tasting nuts, making the trees suitable for commercial or backyard nut orchards. Both kinds of ClINTON 1912 American Chestnut 25 COUrTESy OF BrAD SMITH American chestnut flowers on a tree near Quabban Reservoir in Massachusetts. trees must have resistance to chestnut blight disease and be well adapted to the New England climate (Anagnostakis 1992). There is now interest in developing DNA tests for genetic maps of chestnut trees (http:\/\/www.fagaceae. org\/web\/db\/index), and we are using specific crosses to study the genetics of resistance to diseases as well as to develop timber and orchard chestnut trees. To make these crosses, we put waxed paper bags over female flowers in late June before they are fertile, then put selected pollen on the flowers in July and cover them up again. This allows us to know the parents of the nuts that form. During our breeding program we have found that many hybrids that are the result of crosses between two different species do not form functional pollen. These male-sterile trees produce male catkins with flowers that never bloom. Although this lack of pollen is a nuisance in the breeding program, it is a feature valued by commercial nut growers--they can plant orchards of male-sterile trees with a few pollen-producing trees and have yields of nuts that are very uniform. When it became clear that at least two genes were responsible for resistance to chestnut blight, we began a back-cross breeding program based on the plan of Charles Burnham (Burnham 1988). Asian trees are crossed with American trees, and the hybrids (partially blight resistant) are crossed to American trees again. If there are two resistance genes, one out of four of the progeny from these back-crosses has one copy of both resistance genes, giving it partial resistance. If there are three genes for resistance, one 26 Arnoldia 66\/4 SANDrA ANAGNOSTAkIS A row of twelve-year-old chestnut hybrids selected for timber qualities. out of eight of the progeny will have one copy of all three resistance genes. Trees with partial blight resistance are crossed again to American chestnut trees. This repeated back-crossing increases the percentage of American genes in the hybrids, and selecting for partial resistance insures passage of the resistance genes. A final cross of two trees with partial resistance should result in one of sixteen trees having two copies of two resistance genes (or one of sixty four trees having two copies of three resistance genes), which will make them fully resistant to the chestnut blight fungus. Biological Control of Chestnut Blight Disease In a 1992 Arnoldia article we described viruses, called \"hypoviruses,\" that infect C. parasitica and keep the fungus from killing trees by reducing its virulence (Anagnostakis and Hillman 1992). Since 1972, when CAES imported American Chestnut 27 kEITH kANOTI, MAINE FOrEST SErvICE, BUGWOOD.OrG The densely spiny chestnut bur encloses several nuts, typically three. American chestnut trees in this Hamden, Connecticut, orchard were treated with biocontrol strains from 1978 to 1981, and 15% of the 71 trees survive as the original trunks in spite of the presence of many cankers. Half of the trees continue to be in a repeating cycle of dying back and resprouting. About one third of the trees died back once, resprouted, and the sprouts are still surviving. Percentage of American genes in back-crossed (BC) hybrid chestnut trees. pARENTs 1. American x Japanese 2. American x F1 3. American x BC1 4. American x BC2 5. BC3 x BC3 AMERiCAN GENEs 100% American genes 0% American genes 100% American genes 50% American genes 100% American genes 75% American genes 100% American genes 87.5% American genes 93.8% American genes 93.8% American genes egg + pollen egg + pollen egg + pollen egg + pollen egg + pollen HyBRiD = 50% A F1 = 75% A BC1 = 87.5% A BC2 = 93.8% A BC3 = 93.8% A BC3-F2 SANDrA ANAGNOSTAkIS 28 Arnoldia 66\/4 virus-containing strains of the chestnut blight fungus from Europe, great strides have been made in understanding how these viruses can keep the fungus from killing trees. The genes of three kinds of these (dsrNA) viruses have been sequenced, and the viruses placed in the genus Hypovirus by Bradley Hillman and his collaborators (Hillman et al. 1994). We have studied the movement of both killing and curing strains of the fungus by birds and insects of several kinds (Anagnostakis 1990; Anagnostakis 1995b; Anagnostakis 2001). Although we have introduced hypovirulent strains of the fungus into forest plots, this biological control has not brought about a general recovery of forest chestnuts in Connecticut. However, it has been successful in an orchard of American chestnut trees at the CAES farm in Hamden, Connecticut, where we introduced hypovirulent strains into every canker that we could reach for four years from1978 to 1981. Now, although half of the trees continue to die back from chestnut blight (and sprout, and die back, etc.), about a third that died back once and sprouted now survive and flower even though they are covered with cankers, and about 15% of the trees are the surviving original stems. resistance and all of the native genetic diversity into the future generations. The first generation offspring will be intermediate in resistance, but subsequent generations will produce trees with full resistance. synthesis of Breeding and Biological Control The crosses that have produced blight-resistant trees for timber have, by necessity, used a rather narrow genetic base, even though different trees were used as parents in each generation. At CAES, this has involved crossing and back-crossing both Japanese and Chinese chestnut trees (C. crenata and C. mollissima) with locally adapted American chestnut trees. Our strategy has been to keep native chestnuts alive and flowering by using our biological control agent. This eliminates the need to search for American trees that have survived long enough to flower. It also lets us use populations in specific forest clearings. By planting resistant trees in the forests and treating the native trees with our biocontrol, native trees will survive to naturally cross with the resistant trees and will incorporate blight A basketful of nuts from a hybrid chestnut orchard. Chestnut Trees for the orchard In addition to selecting timber trees, we have continued to evaluate trees for their potential for orchard production in New England. A few acres of chestnut trees can produce enough nuts to sell at farmer's markets or to local stores. The only serious pest is chestnut weevil, which can be controlled by spraying insecticide when the nuts are ripening, or by allowing chickens or guinea fowl to range under the trees and eat the weevils and their grubs. Squirrel control is also essential and every nut farmer has his or her own method. The most productive chestnut orchards are planted with named cultivars, which are vegetatively propagated clones of the original named trees selected for efficient nut production. Since cuttings of chestnut trees will not form roots, chestnut orchard cultivars must be grafted onto suitable rootstock for propagation. Although this increases the cost of the plants, the value in having proven clones makes the purchase price well worth it. Another challenge faced by growers is that some splendid cultivars that do well in one part of the country do not do well in other places. For example, cultivars suited to the far south or to the far west may not do well in New England. Selections from Ohio have generally proven SANDrA ANAGNOSTAkIS American Chestnut 29 JErry A. PAyNE, USDA AGrICUlTUrAl rESEArCH SErvICE, BUGWOOD.OrG reliable in southern New England, as have the few cultivars released from CAES. Since I am the International registrar for Cultivars of Chestnut, information on new trees usually crosses my desk, and I keep a list of the names used and some of their characteristics on our website (http:\/\/www.ct.gov\/caes\/cwp\/view. asp?a=2815&q=376864). The biggest challenge to development of a nut industry in New England is the lack of an established market--many people have never eaten chestnuts and are hesitant even to try them. Also, many who have bought chestnuts and then had weevil larvae crawl out of them will never buy them again. Efforts to develop markets and grower awareness in Michigan and Missouri are making some progress and can serve as examples for New England. The Next problem Even as progress was being made toward blight resistance, another serious chestnut pest arrived. The oriental chestnut gall wasp, Dryocosmus kuriphilus, was introduced into the United States in 1974 by a grower who evaded plant quarantine (Payne et al. 1976). The insect lays its eggs in leaf and flower buds, resulting in defoliated trees with no flowers. Entomologist Jerry Payne chronicled the devastation of orchards of Chinese chestnut trees planted in the state of Georgia. We have reports of infestations throughout Alabama, North Carolina, and Tennessee, and most recently in Columbus, Ohio. As a consequence, breeding work must now include selection for resistance to this pest. Jerry Payne has observed that American and Chinese chinquapins (Castanea pumila, C. ozarkensis, and C. henryi) are resistant to infestation, as are some cultivars of C. crenata. Once again, the CAES collection of species and hybrids is being used for making new crosses, and progeny from these crosses are being tested in North Carolina where the insect is now endemic. These trees were examined by Stacy Clark of the United States Forest Service in 2006 and the preliminary results were encouraging. Of 93 trees planted in 1995, there were 53 that survived the droughts, deer, rabbits, and weed competition for 12 years. Among the survivors, Developing gall and damaged chestnut shoot caused by the oriental chestnut gall wasp. 11 had no wasp galls and 25 had few galls. We hope to understand how resistance is inherited and will incorporate this resistance into our trees as quickly as possible. The other ray of hope for dealing with gall wasp is that Asian parasites released by Jerry Payne seem to be moving with the wasp (Payne et al. 1976). lynne rieske recently reported that parasites were now in the Ohio population (rieske 2007). If these parasites continue to improve as control agents for gall wasp, it is possible that only stressed trees will be seriously damaged by wasp infestation. What's Next? We will soon have timber chestnut trees that can survive in New England. These trees will provide another source of lumber and will also increase the diversity of tree species in forests. We are learning about growing chestnuts in orchards in New England and selecting better FrOM CONNECTICUT ExPErIMENT STATION BUllETIN 154 (1906) This 1905 photograph shows the tall, straight trunk of a then 103-year-old American chestnut in scotland, Connecticut. American Chestnut 31 nut-producing cultivars to make a new niche crop for farmers. The work goes slowly, but is very satisfying. When I talk to scientists who conduct laboratory research, and expect results within months, they are often astonished that I have been working at this research for more than 40 years. There are no quick solutions to the complicated problems in the environment, and trees take a long time to grow. When many factors are interacting they must all be considered. We can make crosses of our trees, wait 10 years for the seedlings to mature, select them, make more crosses, wait 10 years, and still miss some crucial clue in the soil or the weather or animals or insects that will affect our hoped-for outcome. When talking with students I try to emphasize the need for patience, keeping an open mind, and noticing everything. \"Publish or Perish\" and \"More Grant Funding for Survival\" are still driving forces that tempt scientists to focus on small things that can be examined in isolation and written up quickly for scientific journals or granting agencies, but it is important to keep looking at the big picture. References Anagnostakis, S. l. 1990. Improved chestnut tree condition maintained in two Connecticut plots after treatments with hypovirulent strains of the chestnut blight fungus. Forest Science 36: 113124. Anagnostakis, S. l. 1992. Measuring resistance of chestnut trees to chestnut blight. Canadian Journal of Forest Research 22: 568571. Anagnostakis, S. l. 1995a. Effect of the Male Parent on Nut Weight in Chestnut. Annual Report of the Northern Nut Growers Association 86: 124127 Anagnostakis, S. l. 1995b. The Pathogens and Pests of Chestnuts. pp. 125145, In: Advances in Botanical Research, vol. 21, Andrews, J. H. and I. Tommerup, eds., Academic Press, New york. Anagnostakis, S. l., and P. Devin. 1998. lack of Effect of Pollen Parent on Nut Weight of Chestnuts. Annual Report of the Northern Nut Growers Association 89: 1517. Anagnostakis, S. l. 2001. The effect of multiple importations of pests and pathogens on a native tree. Biological Invasions 3: 245254. Anagnostakis, S.l. Chestnut cultivar names. http:\/\/www. ct.gov\/caes\/cwp\/view.asp?a=2815&q=376864 Anagnostakis, S.l. Chestnuts and the introduction of chestnut blight. http:\/\/www.ct.gov\/caes\/cwp\/ view.asp?a=2815&q=376754 Anagnostakis, S. l. and Hillman, B. 1992. Evolution of the chestnut tree and its blight. Arnoldia 52 (2): 210. Burnham, C. r. 1988. The restoration of the American chestnut. American Scientist 76: 478487. Clinton, G. P. 1912. Chestnut bark disease. report of the Station Botanist. 19111912, pp. 407413. Annual Report of The Connecticut Agricultural Experiment Station, New Haven, Connecticut. Cogbill, C. v., Burk, J., and Motzkin, G. 2002. The forests of presettlement New England, USA: spatial and compositional patterns based on town proprietor surveys. Journal of Biogeography 29: 12791304. Graves, A. H. 1937. Breeding new chestnut trees. Annual Report of the Northern Nut Growers Association 28: 93100. Hawes, A. F. 1906. Chestnut in Connecticut and the improvement of the woodlot. Bulletin 154, The Connecticut Agricultural Experiment Station, New Haven, CT. Hillman, B. I., Fulbright, D. W., Nuss, D. l., and van Alfen, N. k. 1994. Hypoviridae. In. Virus Taxonomy: Sixth Report of the International Committee for the Taxonomy of Viruses. F. A. Murphy, C. M. Fauquet, D. H. l. Bishop, S. A. Ghabrial, A. W. Jarvis, G. P. Martelli, M. P. Mayo, and M. D. Summers, eds. Springer-verlag, Wein, New york. Metcalf, H. and J. F.Collins. 1909. The present status of the chestnut bark disease. Bulletin 141, part 5, pp. 4553. U.S. Department of Agriculture, Washington, DC. Payne, J. A., Green, r. A., and lester, D. D. 1976. New nut pest: an oriental chestnut gall wasp in North America. Annual Report of the Northern Nut Growers Association 67: 8386. Pierson, A. H. 1913. Wood-Using Industries of Connecticut. Bulletin 174, The Connecticut Agricultural Experiment Station, New Haven, CT. rieske, l. k. 2007. Success of an exotic gallmaker, Dryocosmus kuriphilus, on chestnut in the USA: an historical account. EPPO Bulletin 37: 172174. Saucier, J. r. 1973. Natural range of American Chestnut, USDA Forest Service Fact Sheet 230. Sandra l. Anagnostakis continues her research at the Connecticut Agricultural Experiment Station in New Haven. She is still having too much fun to consider retiring. "},{"has_event_date":0,"type":"arnoldia","title":"Book Review: American Chestnut: The Life, Death, and Rebirth of a Perfect Tree","article_sequence":4,"start_page":32,"end_page":33,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25457","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14ebb6b.jpg","volume":66,"issue_number":4,"year":2009,"series":null,"season":null,"authors":"Heimarck, Heather D.","article_content":"Book Review: American Chestnut: The Life, Death, and Rebirth of a Perfect Tree Heather D. Heimarck American Chestnut: The Life, Death, and Rebirth of a Perfect Tree Susan Freinkel. University of California Press, 2007. 284 pages. ISBN 978-0-520-24730-7 merican Chestnut: The Life, Death, and Rebirth of a Perfect Tree chronicles the history of chestnut blight, a devastating fungal disease first identified in 1904 by Hermann Merkel, Chief Forester of the New York Zoological Park, and studied by William Murrill, a mycologist at The New York Botanical Garden. The fungus--reclassified in 1978 as Cryphonectria parasitica--swept rapidly through American chestnut's native range, nearly annihilating this once-dominant tree species. By mid-century the blight had reached the southernmost part of the range--Alabama, Mississippi, and northern Georgia. Freinkel notes, \"A map produced by Gravatt in 1943 showed the scope of the pandemic as a long ellipse stretching nearly the full length of the Atlantic seaboard. Within that ellipse, 50 to 99 percent of the chestnuts were dead . . . All told, it is estimated the blight killed between three and four billion trees, enough to fill nine million acres. That is enough trees to cover Yellowstone National Park eighteen hundred times over.\" Susan Freinkel presents ethnobotanical information on the cultivation of the American forests, first by Native Americans and later by rural inhabitants. She describes an Appalachian culture benefiting from an economy based on collecting chestnuts from family \"orchards\"-- actually chestnut stands in the wild that loosely belonged to different families by tradition or proximity. With little else to use as barter, Appalachian families used chestnuts in trade for store goods at mountain exchange posts. The tree's carroty flavored nut was considered superior to other endemic nut species, and its lumber was straight, strong, and rot resistant. The Appalachian voices in this book provide a soliloquy to the species, which was once so abundant that a squirrel could supposedly pass from \"Maine to Georgia\" in its branches. The American chestnut--in diminished form--still lives on more than one hundred years after the blight was first diagnosed. But the species is on a life line, as the author details, waiting for a positive outcome from the experimental strategies of back breeding and guided natural selection. Freinkel conveys the tale of American chestnut through the facets of rural and suburban culture, focusing primarily on Appalachia, New York City, and Pennsylvania. She details the endeavors of the nascent forestry and agricultural departments, observant naturalists and scientists, and well-intentioned legislators. The book portrays key agents and events in the American chestnut's struggle to survive. Many of the chapters are defined by singular people with some intuitive knowledge and skill who bucked common opinion in their methods. They were agents in early control measures like fungal identification, eventual experiments to fight fungus with fungus, and later breeding and scientific efforts to improve the chestnut gene pool. This legacy is mostly borne by a few tenacious individuals, many of whom receive well-deserved public recognition in this book. They were the architects of experimental nurseries and laboratories working on breeding projects or fungal experiments whose results are clocked in a life cycle longer than that of human generations. These efforts, not yet abandoned, may still succeed. This wonderful book is paced like a mystery novel, complete with fascinating characters. The plot line of chestnut's survival includes A Book Review 33 serendipitous interventions such as that of a cross-country skier turned horticulturist who recognized a surviving stand of chestnut trees in Michigan, or the observant tourist who did helpful comparative research on the European chestnut blight. The history also includes unfortunate, foolhardy visions borne of the spirit of the times such as seed irradiation or the advice given to the struggling public to cut down every tree while the lumber value still yielded a profit. The effects of such commerce consequently spread the blight and reduced the gene pool. The story of the American chestnut showcases a chapter of scientific history, human history, and a change in environmental consciousness. Susan Freinkel combines an easy narrative style with a factual yet poetic voice that elevates this material beyond dry science to make it a compelling, addictive read. As the author points out, in a world where a species is lost every minute, the survival and potential comeback of the American chestnut is a victory song for the unsung soldier. The beauty of this book is that at its heart it is a tale of the heroic spirit of individuals who have dedicated careers to work on a solution against great odds. Her observations focus the dialogue on the evolution of a consciousness about an enemy that had not been understood or apprehended. It became an enemy that schooled young foresters, botanists, ecologists, enthusiasts, and scientists on how to work on a problem of vast scope. To the author's credit, she refrains from moralizing or predicting the future. She turns the problem around for proper examination from all sides. An old riddle is answered--yes, if a tree falls in the forest and there is no one there to hear it, it does make a sound. Heather D. Heimarck is Director of the Landscape Institute of the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"2008 Weather at the Arboretum","article_sequence":5,"start_page":34,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25454","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14eb728.jpg","volume":66,"issue_number":4,"year":2009,"series":null,"season":null,"authors":null,"article_content":"2008 Weather at the Arboretum T he autumn of 2007 was so dry that fall transplanting had to be postponed, so moisture was a major concern as 2008 arrived. Fortunately, the year brought greater than normal rainfall which provided optimum conditions for our moisture-starved collections. Arnold Arboretum Weather Station Data 2008 Avg. Max. (F) JAn Feb MAr APr MAy Jun Jul Aug SeP OcT nOv Dec 38.4 39.6 44.2 58.7 65.8 78.4 84.2 75.6 69.2 60.3 59 42.3 Avg. Min. (F) 22.3 23.5 30 37.9 46.8 59.8 65.7 60.8 55.5 40.9 33.2 24.6 Avg. Temp. (F) 30.4 31.6 37.1 48.3 56.3 69.1 75 68.2 62.4 50.6 41.1 33.5 Max. Temp. (F) 65 62 60 72 83 99 94 87 87 74 69 63 Min. Temp. (F) 4 9 11 28 31 51 61 53 42 26 19 8 Precipi- Snowtation fall (inches) (inches) 3.23 8.53 5.65 3.78 1.59 3.97 7.65 5.25 7.24 1.82 4.38 7.84 15 6 14.7 2 January was marked by warmer than normal temperatures, including a balmy 65F on the 8th, and only one snowstorm (on the 14th and 15th) which produced 6 inches of snow, the total for the month. February was mild and wet with over 8.5 inches of total precipitation including 15 inches of snow. Only 2 inches of snow were recorded in March, well below normal, but rainfall added up to nearly 6 inches. April started warm, with some rain, but became dry as the month progressed, causing concern for the imminent spring transplanting season. Our concerns diminished as we received 2 inches of rain toward the end of the month. May was quite dry with only 1.5 inches of rain, and irrigating our new plantings was a priority as we feared a repeat of the dry summer of 2007. Though May's total rainfall was low, a string of days with brief late afternoon or evening showers (which left foliage wet overnight) led to ideal Average Maximum Temperature . . . . . . . . . . . . . 59.6 Average Minimum Temperature . . . . . . . . . . . . . 41.8 Average Temperature . . . . . . . . . . . . . . . . . . . . . . 50.7 Total Precipitation . . . . . . . . . . . . . . . . . . . . . . . . 60.93\" Total Snowfall. . . . . . . . . . . . . . . . . . . . . . . . . . . . 37.7\" Warmest Temperature . . . . . . . . . . . . . . . . . . . . . 99 on June 11 coldest Temperature . . . . . . . . . . . . . . . . . . . . . . 4 on January 4 last Frost Date . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 on May 1 First Frost Date . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 on October 24 growing Season . . . . . . . . . . . . . . . . . . . . . . . . . . 176 days Weather Sue PFeiFFer 35 NANCy rOSe Putnam Fellow Abby Hird installs one of eighteen weather recording units that were placed on the grounds in 2008. Data from these units will be used to create maps of the microclimates within the Arboretum. Julie COOP 2008 was an excellent year for fall color. The foliage of this 105-year-old sweet birch (Betula lenta 17679-A) was in full golden glory on October 29th. Sue PFeiFFer Anthracnose, a fungal disease, was common on the foliage of Fraxinus (seen here), Cornus, Platanus, and Quercus in spring 2008. Heavy rain on frozen ground led to flooding along Willow Path in December 2008. conditions for several fungal diseases including anthracnose. Fears of drought subsided as June was on the wet side with about 4 inches of rain. The temperature on June 11th reached 99F, our high for the year. July was wet and warm, with rainfall and mean temperature above normal. August ended wet and cool with no readings over 90F. These two summer months brought many thunderstorms and buckets of rain--an astonishing total of 12.9 inches for July-August. Flooded roads and streams (and clouds of mosquitoes) were common at the Arboretum and throughout the region. The moist summer allowed us to concentrate on watering only new plantings and not the entire collection, and we did very little supplementary watering overall. September continued our \"above average rainfall\" theme as we transitioned into fall. Heavy rains occurred at the end of the month, totaling about 4 inches in four days. October was sunny, cool, and dry with less than 2 inches of rain--a perfect fall month. The fall foliage display was one of the best the region had seen in many years. The first half of November was mild, but a cold and wet second half transitioned us to what would lie ahead. December was slightly warmer than normal but several major storms piled up a total of 15 inches of snow. Heavy rain (over 2 inches) on the 11th and 12th fell on frozen ground, causing stream flooding and some damage to Arboretum paths; fortunately, no collections plants were damaged. Bob Famiglietti, Horticultural Technologist, compiled the 2008 weather data and wrote the month-by-month weather summary. Additional weather-related information was provided by Stephen Schneider, Manager of Horticulture, and Julie Coop, Manager of Plant Health. "},{"has_event_date":0,"type":"arnoldia","title":"Japanese Clethra: A Hidden Gem","article_sequence":6,"start_page":1,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25458","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14e816f.jpg","volume":66,"issue_number":4,"year":2009,"series":null,"season":null,"authors":"Schulhof, Richard","article_content":"Japanese Clethra: A Hidden Gem A t the edge of the Arnold Arboretum's Central Woods, far from most visitors, grows an exceptional specimen of Japanese clethra (Clethra barbinervis, accession 13087). I first became enamored with this species as a student at Longwood Gardens, where I admired its elegant form as part of the backdrop to the famed Flower Garden Walk. While I later saw several very fine examples of Japanese clethra in the great gardens of the Delaware Valley, none approached the singular beauty and character of the Arnold Arboretum's specimen. Like many plants in the Arboretum's collections, this accession comes with an impressive pedigree, tracing back to Japan in 1886. In that year, William Penn Brooks, a Massachusetts native and valedictorian of the state agricultural college class of 1875, sent seeds of several species to the Arnold Arboretum. Brooks, then a teacher and administrator at Sapporo Agricultural School, found time to survey the surrounding countryside of Hokkaido for interesting plants, several of which came to enrich the Arboretum, including katsura tree (Cercidiphyllum japonicum) and hardy kiwi (Actinidia arguta). Of these plants, the Japanese clethra accession is Brooks's greatest Arboretum legacy. At 122 years of age, the specimen is over 20 feet (6 meters) tall and nearly 15 feet (4.5 meters) wide. In the forests of Japan and Korea, Japanese clethra is said to attain heights of over 30 feet (9 meters), but in North America I have seen few specimens larger than the Arboretum's accession. Although its American cousin, summersweet (Clethra alnifolia), is better known to gardeners, the Arboretum's E. H. Wilson considered C. barbinervis to be the finest ornamental in the genus. Unlike summersweet, Japanese clethra is more a small tree than a shrub. Judicious pruning can emphasize the small-tree form; when trained as such, it displays sinuous single or multiple trunks broken by floating tiers of foliage. This treatment also shows the tree's exquisitely mottled cinnamon, salmon buff, and slate grey bark to full advantage. Japanese clethra's leaves are deep green through summer. Autumn color tends to be unreliable. I have seen outstanding tints of red and burgundy on specimens in both southeastern Pennsylvania and here at the Arnold Arboretum, but in other years have noted little color change before the leaves fall away inconspicuously. Japanese clethra's white flowers are similar to those of summersweet, but they are borne in 4 to 6 inch long panicled racemes that nod forward and gently twist. Japanese clethra blooms around mid July at the Arboretum, about two weeks before our native, and though some consider the fragrance inferior to summersweet, my nose finds it ample in portion and delicious in scent; if anything, too much of a good thing. Clethra barbinervis is listed as hardy to USDA Zone 5 (average annual minimum temperature -10 to -20F [14 to -4C]). Our venerable specimen has survived the coldest of Boston winters, but has proven vulnerable to drought. During a very dry summer in the early 1990s, the death of this specimen seemed certain, with each leaf appearing as if torched to a crisp. Phoenix-like, it recovered, living on as one of the Arboretum's most distinguished, and best-hidden, centenarian plants. Richard Schulhof is Deputy Director of the Arnold Arboretum. RICHARD SCHULHOF Richard Schulhof "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23412","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd160a328.jpg","title":"2009-66-4","volume":66,"issue_number":4,"year":2009,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Impact of Climate Change on the Flora of Thoreau's Concord","article_sequence":1,"start_page":2,"end_page":9,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25452","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14eaf6b.jpg","volume":66,"issue_number":3,"year":2009,"series":null,"season":null,"authors":"Miller-Rushing, Abraham J.; Primack, Richard B.","article_content":"The Impact of Climate Change on the Flora of Thoreau's Concord Abraham J. Miller-Rushing and Richard B. Primack All pHOTOS By THe AUTHOrS UNleSS OTHerWiSe iNDiCATeD C limate change is driving major shifts in ecosystems all over the world, including here in the United States. Tree swallows and many other bird species are breeding earlier, forest edges are extending up the sides of mountains, and the distributions of pest insect species such as the hemlock woolly adelgid are shifting northward. Notably, most of the evidence for biological responses to climate change, including these examples, is based on studies of one or a few species. The number of examples is large, but it is difficult to know how representative they are. How is climate change affecting entire natural communities of plants and animals? Are all of the species within a community changing, or are just a few? Are most species in a location changing in the same way, or are there substantial differences among species? Are there ways to predict how species within communities might change and what the consequences of these changes will be? Despite the obvious importance of these questions, we do not really know the answers. We know many changes are happening, some of them major, but thus far our knowledge is limited to a relatively few species in a few places. To help answer the question of how The statue of Henry David Thoreau at Walden Pond, with a replica of his plant communities are responding to cabin in the background. climate change, we turned to one of also of flowering times. One of these two botathe best-documented floras in the country-- nists was the well-known philosopher and natthe flora of Concord, Massachusetts. The flora uralist Henry David Thoreau, and the other was has been inventoried five times since 1830, a a local shopkeeper, Alfred Hosmer. Between huge effort for the flora of a single town. During them, they recorded the flowering times of over two of these inventories, the botanists collected 700 plant species in Concord. observations not only of plant occurrences, but Flora of Concord 3 affect relationships among plants competing for resources, between plants and herbivores, and between predators and their prey. in the Netherlands, pied flycatcher populations are already declining because they are breeding too late to feed on their caterpillar prey. More broadly, the populations of european migratory birds species that are not migrating earlier in response to warming temperatures are declining, possibly because of temporal mismatches between their migrations and breeding and their environments. Thus, we are left to question, why do species respond differently to climate change? What species are Concord is home to a wide diversity of wildflowers such as this Britton's most (or least) sensitive to changes violet (Viola brittoniana). in climate? Are there characteristics We know of no other climate change study that make a species particularly sensitive? The in the United States that has recorded observaobservations of Thoreau and Hosmer provided tions on as many species in a single location an opportunity to make headway in answering for as long a period as Thoreau and Hosmer. these questions. These observations have major scientific value because we can use them to examine the response of an entire flora to climate change. Their value is further enhanced because changes in the timing of phenological events-- those biological events like flowering, fruiting, and migrations that recur on a seasonal basis--are among the most sensitive biological responses to climate change. Many phenological events in different places in the world are now occurring earlier than in the past. However, it is clear that species' phenologies are changing at different rates. For example, in england some species are flowering more than a month earlier than they did 50 years ago, while other species' flowering times are advancing more slowly, or are not changing at all. in some instances, plants are even flowering later than they did in the past. This variation has the potential to alter important relationships among species, such as those between plants and pollinators. if a plant is flowering much earlier now than it did in the past, but its preferred pollinator is active at the same time each year, the plant and the pollinator could be misThe flowering time of sweet birch (Betula lenta), seen here, matched in time, to the disadvantage of both. proved to be much more responsive to temperature than did the flowering time of gray birch (Betula populifolia). Similarly, variation in changes in timing could 4 Arnoldia 66\/3 The Remarkable History of Botanical Investigation in Concord The surveys of Concord's flora since 1830 are remarkable not only for their number, but also for their intensity. Aside from simply noting species occurrences, in some cases the surveyors recorded species abundance and the presence of invasive species, and noted botanically important areas. Building on the extraordinary historical base of previous surveys of Concord's flora, over the past five years we have conducted our own survey of the flora. All of the previous surveys provided interesting data, but the precise observations of flowering times by Thoreau and Hosmer proved to be the most important for our study. in the 1850s, after a decade of observing nature and four years after his experience of living on How suddenly the flowers bloom! Two or three days ago I could not, or did not, find the leaves of the crowfoot. To-day, not knowing it well, I looked in vain, till at length, in the very warmest nook in the grass above the rocks of the Cliff, I found two bright-yellow blossoms, which betrayed the inconspicuous leaves and all. The spring flowers wait not to perfect their leaves before they expand their blossoms. From the journal of Henry David Thoreau Flora of Concord 5 the edge of Walden pond, Thoreau began recording flowering times of plants in Concord. He cared deeply about the seasonal changes, as can be seen in his many observations in Walden. in describing his activities at Walden, Thoreau states, \"i want to go away soon and live away by the pond . . . But my friends ask what i will do when i get there. Will it not be employment enough to watch the progress of the seasons?\" One might say he was obsessed with the progress of the seasons. From 1852 to 1858, he hiked around Concord and made regular observations of the first flowering times of over 500 different species of The influence of temperature on plant development is shown by lily of the valley plants in an effort to create a (Convallaria majalis). Plants next to a warm building have undergone more develcalendar of the natural events opment than plants further away. in Concord. His intention was to write a book about the seasons in Concord. and Hosmer made was exceptional. They made Unfortunately, he died before he was able to observations across an entire town (without the complete his project. His friends also wrote aid of a car) several days each week for many about his obsession with the seasons, as ellery consecutive years. in the processes, they creChanning did after a walk with Thoreau on ated a list of flowering phenology for more speMarch 6, 1859: \"Our round of walks is as regucies in a single location than any other of which lar as the seasons; now to low spots to look for we are aware. early spring plants, also for early birds. Nature Linking Historical Observations with is an eternal provision and repetition. H[enry] Climate Change Science says there is nothing but the seasons.\" When we learned about these flowering time His efforts to record flowering times were folobservations from the late Thoreau scholar, lowed by another botanical enthusiast, Alfred Brad Dean, and the active New england botaHosmer, who recorded the first flowering times nist, ray Angelo, we recognized that we had an of over 700 species of plants in 1878 and from opportunity to test whether climate change had 1888 to 1902. We do not know exactly why affected the flowering times across the entire Hosmer made these observations, but we are community of flowering plants in Concord. We quite thankful that he did. His records can all immediately set out ourselves to document the be found in precisely organized notebooks, prescurrent flowering times of as many species as ently housed in the Special Collections Section we could throughout the town. From 2003 to of the Concord Free public library. 2006 we visited Concord two to three days each The idea of tracking phenological events was week throughout the flowering season, from not new at the time of Thoreau and Hosmer's March to October, and recorded the plants we work. The practice is said to be as old as agriculsaw in flower each day. We deliberately sought ture, if not older. Well-known Americans such out locations of difficult-to-find species, such as Thomas Jefferson kept records of flowering as Britton's violet (Viola brittoniana) and rose and bird migrations. But the incredible effort pogonia (Pogonia ophioglossoides), with hopes and continuity of the observations that Thoreau 6 Arnoldia 66\/3 of observing as many of Thoreau's and Hosmer's species as we could. eventually, we made observations of the first flowering dates, in addition to recording the entire period of flowering, of over 500 species. Many of the species we observed, and even the places where we saw them, were identical to those that Thoreau and Hosmer had seen. With these data in hand, we set out to test whether first flowering dates had changed in Concord, and whether species differed in their response to a warming climate. To simplify things, we started by analyzing changes in the first flowering dates for 43 of the most common spring-flowering species that Thoreau, Hosmer, and we had all observed in nearly every year we looked. These species were abundant and widely distributed, and probably reflected fairly the changes in flowering times that had occurred more broadly in the Concord flora. On average, these 43 species were indeed flowering about a week earlier in recent years than they had in Thoreau's day, with Hosmer's observations right in the middle. Some of these species' flowering times changed dramatically. For example, highbush blueberry (Vaccinium corymbosum), a shrub of wetlands, and yellow wood sorrel (Oxalis stricta), a native herb of fields and roadsides, are now flowering 21 and 32 days earlier, respectively, than they did 150 years ago. This trend toward earlier flowering corresponded with warming temperatures in the Concord area. Temperature records from the Blue Hill Meteorological Observatory in Milton, Massachusetts showed 2.4C (4.3F) warming from 1852 to 2006. Most of this warming occurred because of urbanization and development in the greater Boston area, while some of it occurred because of global climate change. (For comparison, the average global temperature warmed by about 0.7C (1.3F) over the past 100 years.) The relatively large warming in Concord, boosted by urbanization, makes this flora a good example for how floras in the rest of the country, away from cities, may respond to future warming. Global temperatures are predicted to warm by about 3C (5.4F) in the next 100 years. Of course, the responses of plants in a developed area may be different from rural areas for a variety of reasons, pollution and the rate of warming among the most important. However, pollution is thought to have a negligible effect on flowering times relative to temperature. And although the rate of warming could alter how quickly flowering responses to temperature might evolve, evolution is likely to be slow in a flora, such as Concord's, dominated by longlived perennials and should have little impact on flowering responses to temperature on the time scales with which we were working. We found a strong relationship between temperature and flowering times in Concord; temperatures in January, April, and May explained most of the variation in flowering dates. (More on why those months later.) On average, plants flowered about three days earlier for each 1C warming. When we examined a much larger list of 296 species, we found the same response of flowering dates to temperatures. Because plant flowering dates respond to temperatures at particular times, we examined the relative importance of temperatures in each month. Temperatures in January, April, and May were by far the most important for most species; warmer temperatures in all three months were linked to earlier flowering dates. April and May were important because they were the temperatures immediately preceding flowering for most species. Why January temperatures were so important was more of a mystery. Winter temperatures are typically understood to influence flowering times in temperate plants through a process known as vernalization. plants have biochemical methods to keep track of how cold it is and for how long. Once it has been cold enough for long enough, plants are said to be competent--that is, their dormancy is nearly complete and they are ready to start developing leaves and flowers as soon as it warms up. if it never gets cold enough for long enough, the plants may still flower when temperatures warm in the spring, but it will take longer to flower; the colder it gets, the closer the plants come to being fully competent, the faster they flower in the spring. The need to track the cold is a defense against abnormally warm temperatures in mid-winter. plants in New england are adapted to avoid flowering in January or February, even when the weather is Flora of Concord 7 Rose pogonia (Pogonia ophioglossoides) is an elusive native orchid found in fens and other damp sites. Cardinal flower (Lobelia cardinalis) grows in moist soils and produces spikes of bright red flowers in mid to late summer. warm. Under this scenario, plants should flower earlier in the spring after cold winters, because they should become more fully competent. However, we found that plants flowered later after particularly cold Januaries. Vernalization was apparently not responsible for the response we saw. instead, we suspect that it gets so cold in Concord in January that in the coldest years plants suffer physical damage to the vascular tissue that delays flowering. We found support for this hypothesis in a subsequent project of ours in which we looked for this sort of damage in birches at the Arnold Arboretum. Variation Among Species With data on all these species, we looked for patterns of variation. Which species were the most sensitive to temperatures? Did particular traits seem to be associated with strong responses to temperature? The flowering responses of plants from different habitats (e.g., aquatic, forest, grassland, roadside, and wetland) did not differ, nor did the responses of natives and non-natives differ from each other. However, season of flowering seemed to matter a great deal. The flowering times of early-season plants were on average strongly correlated with temperature, whereas the flowering times of late-season plants had a much weaker link with temperature. if this pattern continues, the flowering season may lengthen as temperatures warm. Spring species may flower ever earlier and the flowering times of summer species may not change much at all. As a result, the degree of overlap in species' flowering times may be reduced. Growth form seemed also to matter, although in a more indirect way. Among perennial herbs, the flowering dates of late-season plants varied a lot from year to year compared to the flowering dates of early-season plants. However, the variation in flowering time was not related to temperature. We are still unsure of what is driving the year-to-year variation in the flowering dates of late-season perennial herbs; it might have do with soil moisture, degree of shading, 8 Arnoldia 66\/3 or land use. Meanwhile, woody plants showed the pattern we expected--lots of climate-driven variation in flowering dates for early-season plants and relative stability in the flowering of lateseason plants. lastly, we noted that the flowering responses of several closely related species varied substantially. For example, sweet birch (Betula lenta) and gray birch (Betula populifolia), which occur in many of the same habitats in Concord and elsewhere, showed very different responses to temperature. Sweet birch flowered about three days earlier for each 1C increase in January, March, and April temperatures, whereas gray birch flowering dates were Bloodroot (Sanguinaria canadensis) is one of the early spring ephemerals recorded unrelated to temperature. in an in Concord's flora. even more dramatic example, ecologically, these results reflect the comrough-stemmed goldenrod (Solidago rugosa) plexity of plant response to climate change. flowered 11 days earlier for each 1C increase The flowering times of early-season plants are in temperature, whereas the flowering dates of shifting more quickly than those of late-sealance-leaved goldenrod (Solidago graminifolia) son plants. perennial herbs and woody plants and most other goldenrods were unrelated to respond differently. Habitat and nativeness do temperature. These varied responses to clinot seem to affect flowering responses to climate change could lead to increased hybridizamate change. With such a wealth of data from tion among closely related species, if flowering this single location, we found surprising pattimes that previously occurred at distinct times terns, leading to questions that still must be began to overlap. it could also cause more comanswered. For example, why do the flowering petition for pollinators; if plants that share times of closely related species respond so difpollinators and have historically flowered at ferently to warming temperatures? How will different times begin flowering at the same species interactions change as a result? Given time, they may start competing for the pollinathe remaining uncertainty, it is difficult to tors' services. Competition for nutrients and assess how exactly flowering times will shift water needed at critical times in plant developin the future and what the changes will mean ment could also increase. for plants and animals. Thoreau was also aware The Big Picture of the effects of climate on plants and animals Our study found that the plant community and their interactions more than 150 years ago in Concord is responding to climate change when he wrote, \"Vegetation starts when the in dramatic ways. Spring is coming earlier on earth's axis is sufficiently inclined; i.e. it folaverage, but there is a lot of variation among lows the sun. insects and the smaller animals species. Highbush blueberries are flowering (as well as many larger) follow vegetation . . . three weeks earlier than they were in ThoThe greater or less abundance of food deterreau's day, yet the flowering times of other mines migrations. if the buds are deceived and species are not changing at all. suffer from frost, then are the birds.\" Flora of Concord 9 The results of our study show how perceptive he was and suggest likely impacts of a warming climate. interactions among species--e.g., predators and prey, competing plants, plants and herbivores--will certainly change. Most plant species can be pollinated by many animal species or by the wind, and most pollinators can feed on various flowers. For them, the relationships will shift, but they will probably continue to persist. However, shifts in the timing of specialist interactions, like those between hummingbirds and plants with long flower tubes, will probably lead to more dire consequences for the species involved. if a plant with a specialist pollinator flowers before its pollinator is active, its chances of reproducing may decline significantly. Additionally, as the flowering times of early-season plants continue to advance and pull away from the flowering times of late-season plants, there may be times of low floral resources for pollinators like bumblebees. Undoubtedly, some species will do well and thrive under the changed circumstances, while others will do poorly and may even go extinct. Thoreau was keenly aware of the importance of educating people about environmental issues. He helped Concord's citizens to appreciate wild nature, and he encouraged them to protect it. He wrote, \"i think that each town should have a park, or rather a primitive forest of five hundred or a thousand acres, either in one body or several, where a stick should never be cut for fuel, nor for the navy, nor to make wagons, but stand and decay for higher uses-- a common possession forever, for instruction and recreation.\" residents of Concord and the government have followed this advice; about 40% of Concord's land is preserved in parks and protected areas, such as Walden pond State reservation, Great Meadows National Wildlife refuge, and the estabrook Woods. With the help of these protected areas, we have been able to continue the same observations of flowering times made by Thoreau at many of the same localities in Concord. We now hope that Thoreau's observations and our own work will promote broad discussion of the effects of climate change on biological systems. Resources: Bradley, N. l., A. C. leopold, J. ross, and W. Huffaker. 1999. phenological changes reflect climate change in Wisconsin. Proceedings of the National Academy of Sciences of the United States of America 96: 97019704. Miller-rushing, A. J., and r. B. primack. 2008. Global warming and flowering times in Thoreau's Concord: a community perspective. Ecology 89: 332341. Miller-rushing, A. J., r. B. primack, D. primack, and S. Mukunda. 2006. photographs and herbarium specimens as tools to document phenological changes in response to global war ming. American Journal of Botany 93: 16671674. parmesan, C. 2007. influences of species, latitudes and methodologies on estimates of phenological response to global warming. Global Change Biology 13: 18601872. USA National phenology Network. www.usanpn.org Abraham Miller-rushing and richard primack completed this research together while working at Boston University. Dr. Miller-rushing is now the coordinator of the Wildlife phenology program, a collaboration between the USA National phenology Network and The Wildlife Society. Dr. primack is a professor at Boston University. What's Next? research efforts are underway to solve these unanswered questions, but more data are needed. Known sets of phenological observations like those of Thoreau and Hosmer are quite rare. yet scores of people have recorded observations--flowering dates in gardens, birds' spring arrivals at feeders--that now sit in boxes in attics and basements. The newly formed USA National phenology Network (www. usanpn.org) is beginning to collect these valuable \"shoebox\" data sets to make them freely available to the research community and the public. There is great potential for these phenological observations to shed light on ecological responses to climate change. in addition, evidence of changes in phenological events can improve public awareness of the effects that climate change is already having on biological systems. people can see changes in phenology in their back yards, neighborhoods, parks, and forests. We believe that building on the observations of a well-known figure such as Thoreau can further increase the potential for public outreach. "},{"has_event_date":0,"type":"arnoldia","title":"A Matter of Taste: Pleasure Gardens and Civic Life","article_sequence":2,"start_page":10,"end_page":14,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25449","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14eab28.jpg","volume":66,"issue_number":3,"year":2009,"series":null,"season":null,"authors":"Andersen, Phyllis","article_content":"A Matter of Taste: Pleasure Gardens and Civic Life Phyllis Andersen \"To be natural is such a very difficult pose to keep up.\" --Oscar Wilde, An Ideal Husband \"P opular taste is not a criterion that those who serve our public can respect.\" So said Mariana Van Rensselaer, the distinguished New York art critic and first biographer of architect H.H. Richardson. That remark, made in 1888, fueled the controversy that erupted over her criticism of flowerbeds in Boston's Public Garden. Describing them as crude hues in false situations, she took particular offense at `Crystal Palace Gem' geraniums: \"The cherry colored blossoms with yellow-green leaves are the most hideous products of recent horticulture.\" William Doogue, the Irish-born horticulturist in charge of the Garden's plantings, took exception to her criticism and also rebuked her social position, personal gardening habits, and Harvard-connected friends. Doogue defended his work as accommodating the general taste of the public, who loved his plantings. He protested to the local newspapers and the Mayor, and anyone else who would hear him out. Was all of this brouhaha caused by some ill-placed geraniums, or was it indicative of a deeper division in how we imagine our public parks? This division is illustrated by the wellknown story of the 1858 design competition for New York's Central Park, won by Frederick Law Olmsted and architect Calvert Vaux with a plan titled \"Greensward.\" Their proposal offered a picturesque landscape evocative of the English countryside, combining rustic structures with meadows punctuated by groves, rock outcroppings, and sinuous water bodies. \"Sylvan\" and \"verdant\" were words used by the designers to describe their design as \"a constant suggestion to the imagination of an unlimited range of rural conditions.\" The contrast with the majority of proposals from competitors-- A source of color and controversy, `Crystal Palace Gem' geranium. engineers, landscape gardeners, and talented amateurs--represented a remarkable shift toward the narrative of the picturesque. Other more traditional plans presented highly embellished gardens with formal promenades, fountains, arches, statues of Greek deities and New York politicians, bandstands, and extensive formal layouts of flowering plants. By the mid nineteenth century, the educated public understood that the picturesque landscape was the aesthetic ideal for public parks, allowing the mind to wander along with the body. Among others whose opinions counted, economist and social critic Thorstein Veblen pointed to an upper-class predilection for public parks that were rustic and natural. Enlightened park advocates rejected the pleasure garden model with its emphasis on flowery display, theatricality, sociability, and amusement, believing its artificiality and \"claptrap and gewgaw\" lacked moral uplift and tasteful restraint. Like sin and grace, the picturesque park and the pleasure garden are mutually defining. Olmsted used medical metaphors to promote his COuRTESY OF JIMMY TuRNER. Pleasure Gardens 11 notion of the park ideal: parks should be an antidote to urban ills, healing places for damaged minds. Calvert Vaux's famous comment on Americans' intuitive love of the country was at the core of learned park discussions. Vaux spoke of an \"innate homage to the natural in contradistinction to the artificial, a preference for the works of God to the works of man.\" Supporters of the pleasure garden model rejected the imposition of rural scenery on the city and embraced the seductive lure of sensual sound, color, and light--a sustained Fourth of July celebration, an extended summer fete. The Origin of the Public Pleasure Garden The public pleasure garden originated in London in the eighteenth century with extensive public gardens established at Ranelagh, Marylebone, and Islington. But Vauxhall Gardens on London's South Bank most completely and intensely captured the public's imagination. A favorite watering hole for Samuel Johnson, it was frequently used as a fictional backdrop by novelists. It offered grand promenades, open-air temples imitating ancient buildings, an array of dining and drinking pavilions, small theatres, bandstands, tea gardens, and private bowers for romantic interludes. Linking the attractions were elaborate flower displays of local and foreign blooms selected for color, fragrance, and mood-evoking exotic origins. There were fireworks and beguiling night-lighting in an era when both were rare. In its heyday, Vauxhall Gardens attracted aristocracy, royalty, and anyone who wished to mingle and immerse in an environment designed to please. New York entrepreneurs transported the Vauxhall Gardens concept, name, and menu of PHYLLIS ANDERSEN Central Park's Sheep Meadow reflects the pastoral, naturalistic theme inherent in Olmsted and Vaux's winning design for the park. 12 Arnoldia 66\/3 Music, dining, and assorted other revelries made London's Vauxhall Gardens the place to see and be seen. Vauxhall Gardens, 1785, engraved by Robert Pollard II after Thomas Rowlandson. credit: The Metropolitan Museum of Art, The Elisha Whittelsey Collection, The Elisha Whittelsey Fund, 1959 (59.333.975). Image The Metropolitan Museum of Art attractions to New York in 1805, to the area around Broadway and East 8th Street, which is now known as Astor Place. At the same time, even the less than sybaritic Hoboken, New Jersey created Elysian Fields, a popular waterfront park that offered ferry service from Manhattan, and where, some say, the first organized game of baseball took place. The last of the New York pleasure gardens, Palace Gardens, opened in 1858 (the same year as the Central Park competition). It offered the usual array of dining pavilions, water features, and elaborate night-lighting. Legacy of the Pleasure Garden Today, the tradition of the pleasure garden continues to influence the way we think about urban parks. Certainly the questions posed 150 years ago continue to resonate: Who owns the parks? The planners? The middle class? The working class having no other options? And just as important: What is the purpose of a park? The success of the public pleasure gardens was due to diligent management by entrepreneurs who owned them and developed new attractions: balloon launches, water gondolas, music commissioned for special occasions. The eventual demise of the public pleasure garden was due in part to competition from new urban amenities: restaurants, concert halls, theatres, tearooms, and cafes dispersed throughout the city. It was due as well to the growth of petty crime that, then as now, often attaches to public venues that draw huge crowds. And some plea- Pleasure Gardens 13 sure gardens, having contributed to the growth and desirability of the city, became victims of their own success and were lost to real-estate development pressures. The prototypical evocation of a pleasure garden that survived is Copenhagen's Tivoli, which opened in 1843. Patterned on London's Vauxhall and named for the beautiful resort town near Rome, it still offers families a complete pleasure garden experience with attractions interspersed among flower displays appropriate to the season. The horticultural display of pleasure gardens, with its emphasis on seasonal flowering, evolved into civic horticulture--embellishment of city-spaces that are not within the purview of the professional landscape architect and most often maintained by gardeners trained through apprenticeship and guided by trade magazines. These plantings typically feature massing of large numbers of flowers of strong color con- trasts arranged in geometric or pictorial patterns. Some traditions, such as the theatrical display of plants in graduated tiers, evolved from the eighteenth-century English estate garden into the public pleasure garden, as still seen in Boston's Public Garden today. Civic horticulture draws on a rich planting tradition that evokes admiration of both the beauty of the plantings and the ingenuity of the gardener. The immense popularity of the Rose Garden in the Fens section of Boston's Emerald Necklace, of the planted borders in downtown Boston's Post Office Square, and the grand flowerbeds at Copley Square are fine examples of horticulture that enlivens the city, akin to Pop Concerts on the Esplanade. Although theme parks and amusement parks are obvious descendents of the pleasure garden, recent trends in urban public parks suggest that the pleasure garden is enjoying a renaissance PHOTOS COuRTESY OF TIVOLI Modeled on public pleasure gardens such as Vauxhall, Tivoli opened in Copenhagen, Denmark, in 1843. Tivoli's exotic Moorishstyled Nimb building is shown in 1910 (left), one year after being built, and as it appears today (right). 14 Arnoldia 66\/3 wood have developed a highly ornamental planting plan for the North End Park of Boston's Rose Kennedy Greenway. The Dutch horticulturist Piet Oudolf is acting as a consultant for a number of new urban parks in the united States, bringing his skill at highly textured perennial planting in changing seasonal patterns to a new audience. Yet, we still drag issues of public taste behind us, although now couched in concerns for environmental suitability, often with the same moral overtones that characterize the Central Park discussions of the midnineteenth century. We lay a huge responsibility on our urban parks. They must be didactic, educate about ecology, unify communities, and convey history. They must exhibit good taste and local values. But if we are to sustain parks in cities, they must embrace the imagination of the public. The term \"Disneyfication\" is now Beds of brightly colored annual flowers feature prominently in views of Boston's Public Garden from an early-1900s postcard (top) and a 2006 photograph (bottom). an indictment, but one suspects that William of sorts. We are in the midst of defining a new Doogue would have welcomed Walt Disney's words: \"We are not trying to entertain critics. urban park discourse, one that rejects the picI'll take my chances with the public.\" turesque and encourages new kinds of urban engagement--drawing in the city, making use of technology, and embracing theatricality. ChiPhyllis Andersen is a landscape historian and the former cago's Millennium Park, an assemblage of culdirector of the Institute for Cultural Landscape Studies tural attractions and elaborate planting displays, of the Arnold Arboretum. She is currently working lists \"theatre consultant and lighting designer\" on a book on public pleasure gardens scheduled for as part of the design team. The team of Kathryn publication in 2010. This article originally appeared in ArchitectureBoston. Gustafson and Crosby, Schlessinger and Small- PHYLLIS ANDERSEN "},{"has_event_date":0,"type":"arnoldia","title":"The Cathay Silver Fir: Its Discovery and Journey Out of China","article_sequence":3,"start_page":15,"end_page":25,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25451","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14eaf26.jpg","volume":66,"issue_number":3,"year":2009,"series":null,"season":null,"authors":"Callaghan, Christopher B.; McNamara, William; Del Tredici, Peter","article_content":"The Cathay Silver Fir: Its Discovery and Journey Out of China AUSTrAlIAn BICEnTEnnIAl ArBorETUM Christopher B. Callaghan ll that glitters isn't gold. Sometimes it's silver, especially when it's the rare Cathay silver fir (Cathaya argyrophylla). It is now over 50 years since the discovery of this \"living fossil\", yet it remains largely unknown. Access to this conifer has been tightly controlled by China; reportedly, even the offer of a Trident jet in exchange for a single plant during the late 1970s was not sufficient to entice the Chinese to release their grip on this endemic \"treasure tree,\" whose fir-like leaves reflect their silvery undersides when they catch the sunlight. This offer may not sound so farfetched when compared with the more recently discovered Wollemi pine (Wollemia nobilis) of Australia, which has grossed millions of dollars in worldProminent stomatic bands on the undersides of leaves give Cathaya argywide sales since its public release in rophylla its silvery flash. early 2007 following a well- orcheswestern cultivation from China in late 1947, trated marketing and publicity campaign higha mere six years after its discovery. This was lighting its ancient origins. Earlier, a pre-release thanks in large part to the efforts of Elmer D. auction of the first Wollemi pines realized over Merrill, then Arnold Professor of Botany at one million Australian dollars with an average the Arnold Arboretum and previously the of A$3,627 per tree, and this without any AmerArboretum's director. ican bids because of U.S. import restrictions on By the time of the discovery of Cathaya argytrees over 18 inches (0.5m) tall. So for China, rophylla just eight years later, the changing the Cathay silver fir--mass produced and proppolitical landscape in China and the cutting of erly marketed to the west--had the potential of ties with the west meant that this botanically being a similar financial success story. interesting tree, which Chinese botanists have Described as another \"living fossil\" when it described as \"The Giant Panda of the Plant Kingmade world headlines in the 1950s, the Cathay dom,\" was to languish in near obscurity for over silver fir did not make it out of China prior to thirty years. Even with the gradual lifting of the its official release by Chinese authorities in bamboo curtain post-1972, it still took many the 1990s. In contrast, dawn redwood (Metaseyears before the Chinese allowed the tree to be quoia glyptostroboides), the previous worldtaken out of the country, or distributed any seeds renowned \"living fossil\", was introduced into A 16 Arnoldia 66\/3 to overseas botanical institutions. Consequently, the Cathay silver fir is still little known even today, more than half a century after its scientific discovery. A New Plant is Found This discovery occurred in 1955 during a botanical exploration of the remote Huaping region of northern Kwangsi province (now Guangxi Zhuang Autonomous region) in southern China. Deng Xianfu, a member of the Kwangfulingchu Expedition, literally unearthed the first Cathay silver fir when he dug up a seedling of what he thought was Keteleeria fortunei. Following a closer inspection of the seedling, expedition leader Professor Zhong Jixin found that it didn't resemble Fortune's keteleeria. He also knew that Keteleeria fortunei, while occurring naturally in Kwangsi province, could not survive there at above 1400 meters (4600 feet) in the Tianping Mountains, and so considered that it might be a new species of Keteleeria. Upon receiving further information that a tree had been seen in these mountains with some resemblance to both a pine (Pinus) and a fir (Abies), Professor Zhong realized that they should be looking for something special. He directed expedition members to intensify their efforts to find the parent plant(s) of the unfamiliar seedling. Continued searching of the precipitous, mist-shrouded mountains led to the discovery of a mature tree on the southern slopes of Mt. Hongya on May 16, 1955. Herbarium specimens were collected by expedition members, with further specimens collected from the same locality by H.C. lei, H.C. Chung, H.l. Hsu and H.F. Tan from May to July the following year. All these specimens were deposited at the herbarium of the South-China Institute of Botany, Sketch maps not to scale Numbers on the enlarged map indicate the approximate locations of wild Cathaya (see facing page). Additionally, Cathaya argyrophylla is said to occur at Luohandong in Hunan province and in Tongzi county, Guizhou province. The approximate position of these two localities has not been determined and is therefore not indicated on the map. C.M. = Chongqing Municipality. later renamed Kwangtung Institute of Botany (now held at Guangxi Institute of Biology). Here they were seen by the Soviet botanist Sugatchey [likely a mistranslation of the name Sukachev] who advised that they resembled plant fossils previously found in the Soviet Union and Europe dating back to the Pliocene of the Tertiary Period, and hence the newly discovered tree represented a \"living fossil\". Cathaya fossils found since then include fossil pollen in Asia and north America dating back to the Cretaceous. Chun Woon Young (Chen Huanyong) and Kuang Ko Zen (Kuang Keren) published a description of the new genus and species in 1958. They also described a second species, Cathaya nanchuanensis, discovered in 1955 on Jinfo Shan (Golden Buddha Mountain) in southeastern Sichuan. However, this name was reduced to a synonym of Cathaya argyrophylla in 1978. Cathay Silver Fir 17 Natural Occurrences of Cathaya in China ProviNCe\/ reGioN GuANGXi ZHuANG AuToNoMouS reGioN NuMber oN MAP LoCATioN (reserve area in ha) Dayao Mountain Nature reserve (aka Dayao Shan National Forest Park) established 1982. Jinxiu County. Huaping Nature reserve (aka Huaping Primeval Forest), Mt Tianping. established 1961. Sanmen, Longsheng County (type specimen of Cathaya argyrophylla found here in 1955 near Yezhutang, Southern slope of Mount Hongya). Cathay Silver Fir Nature reserve (aka Dashahe Cathaya reserve) established 1984. Daozhen Xian, Daozhen County. Forest reserve of Guizhou botanical Garden. Founded 1964. Liuchongguan, Guiyang. Mount Fanjing Nature reserve. established 1978. Jiangkou County. Dingliao Nature reserve. established 1986. Zixing County\/ bamian Mountain Nature reserve est. 1982. Guidong County. Ziyunwanfeng Mountain Nature reserve. established 1982. Xinning and Chengbu Counties. Wulong County. Jinfo Mountains Nature reserve. established 1979. Nanchuan County. (Cathaya nanchuanensis, now regarded as an ecotype of C. argyrophylla, found here in 1955.) GuiZHou HuNAN CHoNGQiNG MuNiCiPALiTY (previously part of Sichuan Province) The generic name Cathaya derives from the historic place name Cathay, a dominion of the Mongol Emperor Kublai Khan at the time of Marco Polo's travels during the late thirteenth century, and now the northern section of today's China. However, the areas of the present day natural occurrence of Cathaya are actually outside the realm of what was known as Cathay in Marco Polo's time. Instead, they fall within another of Kublai Khan's dominions known as Mangi or Manzi, now the region of China south of the Yangtze river. So perhaps in a historical context the name Mangia would have been more appropriate, although without the appeal of implied antiquity in the name Cathaya. Guarding the Silver The significance of the discovery of the Cathay silver fir in 1955 was considered by the Chinese to be so important that they established Hua- ping nature reserve in 1961 to protect the first found population of the trees. This was one of the earliest nature reserves created in China. Since 1976 many more nature reserves have been established throughout China, and around 4,000 Cathay silver firs presently occur in about a dozen of these (see map). Even when China opened to the west in the late 1970s, these nature reserves were generally off-limits to most foreigners. As late as 1997, I and my colleague S.K. Png of the Australian Bicentennial Arboretum, during a visit to Guizhou Botanical Garden in Guiyang, were steered clear of the natural stand of Cathaya argyrophylla growing in the forest reserve of the garden. A similar situation befell the authors of Southwest China, Off the Beaten Track while they were researching their book during the mid-1980s, and were discouraged from visiting Huaping nature Preserve in longsheng County, 18 Arnoldia 66\/3 AUSTrAlIAn BICEnTEnnIAl ArBorETUM A miniature forest of Cathaya argyrophylla seedlings emerges from a propagation flat. where Cathaya argyrophylla was first found. As they commented, \"longsheng County has a nature preserve, though what is there is anyone's guess since we could never get a straight answer\". [Ed. note: William Mcnamara of Quarryhill Botanical Garden also had a challenging experience trying to see Cathaya--read his account on page 24] Ultimately, China must have realized that one way to protect these rare and endangered trees in their natural habitat is to make them A juvenile plant of Cathaya argyrophylla growing at the Australian bicentennial Arboretum. AUSTrAlIAn BICEnTEnnIAl ArBorETUM Seven- and ten-year-old trees at the Australian bicentennial Arboretum are the first known Cathaya in cultivation outside of China to bear male and female strobili (a male strobus on the seven-year-old tree is shown here). in China, cultivated specimens are said to take as long as 17 years to bear male flowers and even longer to bear fruit. available for cultivation elsewhere. The earliest record I've found for Cathaya argyrophylla introduced outside China is a 1993 accession at the royal Botanic Gardens, Sydney, Australia, The accession's exact fate wasn't recorded, but as of May, 2003, it was listed as \"no longer in the nursery\". However, since plants which had lost their identification labels in the botanic garden's nursery were sometimes sold at the annual Friends of the Garden's plant sales, it is at least possible that the oldest Cathaya in cultivation outside of China is growing unrecognized in a yard somewhere in Sydney. The next earliest year for introduction of definitely surviving Cathaya argyrophylla is 1995 when seeds were received by the royal Botanic Garden, Edinburgh, Scotland from Shenzhen Botanical Garden in China. These seeds were then redistributed by Edinburgh's Conifer Conservation Program to various other gardens, AUSTrAlIAn BICEnTEnnIAl ArBorETUM Cathay Silver Fir 19 DEnDroloGICAl ATlAS ProJECT TEAM--ZSolT DEBrECZY, ISTvan raCZ A native stand of Cathaya argyrophylla grows on a steep, misty mountainside in China. including 50 seed sent to the Arnold Arboretum, where none germinated. The Arnold Arboretum received further seed in 1998 from Fairy lake Botanical Garden in China, with excellent germination [Ed. note: read more about Cathaya at the Arnold Arboretum on page 22]. Finally, seed was allowed out of China in commercial quantities in 1998. Worldwide, apart from botanical gardens, arboreta, and rare plant collectors, relatively few private individuals appear to have acquired this desirable conifer, although many have expressed interest in obtaining the plant if and when it becomes available. Cathaya in the landscape Although not yet widely grown, Cathay silver fir certainly has potential as a landscape plant. It is beautiful as a young plant, and ultimately develops into a noble tree of about 20 meters (65 feet) or more tall with a columnar trunk and horizontal branching. Its long, narrow, evergreen leaves are about 4 to 6 centimeters (1.4 to 2.4 inches) long (sometimes longer), and 2.5 to 3 millimeters (.08 to .11 inches) wide. leaf color is deep green. on the underside, two prominent silvery-white stomatal bands are separated by the midrib. This flash of silver provides the species with its specific epithet, argyrophylla, \"with silvery leaves\". Surviving as it does in Chinese botanical gardens at Shanghai near the coast and Wuhan in central China, which experience minimum winter temperatures of -12C ( 10F) and -18C (0F) respectively, this rare and endangered tree should be hardy in USDA zones 7 or warmer. In slightly colder regions it may be suited to cultivation provided it is given a sheltered microclimate where it is protected from extremes of winter cold and freezing winds. In its native range Cathay silver fir experiences cool summers, winter snow, high humidity, and plen- 20 Arnoldia 66\/3 C Two Living Fossils and the Arnold Arboretum Connection athaya argyrophylla co-author Chun Woon Young (Chen Huanyong) had undertaken dendrology courses with Professor John Jack at the Arnold Arboretum from 1915 to 1919 while completing graduate studies at Harvard's Bussey Institution. He was to comment that it would take him a lifetime of travel to learn as much about Chinese trees as he did while studying at the Arnold Arboretum for a few years. Hsen Hsu Hu (Hu Xiansu), who was the lead author with W.C. Cheng in naming and describing the dawn redwood (Metasequoia glyptostroboides), also studied under John Jack from 1923 to 1925. Thus, both the monotypic \"living fossil\" conifers endemic to China, Cathaya argyrophylla and Metasequoia glyptostroboides, were named and described by pioneering Chinese botanists who undertook forestry courses at the Arnold Arboretum. ArCHIvES oF THE ArnolD ArBorETUM in this 1917 photograph, Professor John G. Jack studies a black maple with several students, including Chun Woon Young (Chen Huanyong) at right. Cathay Silver Fir 21 DEnDroloGICAl ATlAS ProJECT TEAM--ZSolT DEBrECZY, ISTvan raCZ tiful rainfall. When planted in the landscape it should grow best if it receives plenty of moisture, particularly in summer, and is situated in a sunny, well-drained site. The Cathay silver fir is one of the most notable in a long line of rare, endemic, and endangered plants to come out of China, Ernest Wilson's \"Mother of Gardens,\" and I suspect that there remain others yet to be discovered. We can only hope that they are found before human population pressure and the resultant clearing of ever-diminishing forested areas forces them to extinction, as is sadly happening throughout the world. references and Further reading Bean, W. J. 1970. Cathaya. In: Trees & Shrubs Hardy in the British Isles (8th Ed.) vol. 1. p. 539. John Murray, london. Callaghan, C. 2006. Botanical Sale of the Century. Wollemi Pine Auction. International Dendrology Society Ye a r b o o k 2 0 0 5 , p p . 4 7 4 9 . Dendrology Charitable Company, Herefordshire, England. Callaghan, C. 2007. A synopsis of the enigmatic Cathay Silver Fir. International Dendrology Society Yearbook 2006, pp. 151164. Dendrology Charitable Company, Herefordshire, England. Callaghan, C. 2007. Bibliography of Cathaya--living & Fossil. International Dendrology Society Yearbook 2006, pp. 164167. Dendrology Charitable Company, Herefordshire, England. A mature Cathay silver fir displays its picturesque habit. Dallimore, W. and A. B. Jackson. 1966. Cathaya. In: A Handbook of Coniferae and Ginkgoaceae. (4th Ed.), pp. 136137. Edward Arnold, london. den ouden, P. and B. K. Boom. 1965. Cathaya. In: Manual of Cultivated Conifers Hardy in the Cold and Warm Temperate Zone. pp. 5354. Martinus nijhoff, The Hague. Farjon, A. 1990. Cathaya. In: Pinaceae: Drawings and descriptions of the genera. regnum veg. 121, pp. 171175. Koeltz Scientific Books, Konigstein. (continues on page 25) Cheng, W. C. and l. K. Fu. 1978. Cathaya. In: Flora R e p u b l i c a e P o p u l a r i s S i n i c a e , vo l . 7 , Gymnospermae. p. 122, plate 30. Science Press, Beijing. [in Chinese]. Chun, W. Y. and K. Z. Kuang. 1958. Genus novum Pinacearum ex Sina australi et occidentali. (A new genus of Pinaceae--Cathaya, from the South and West of China). Bot. Zhurn., Moscow, 43(4):461476, 10 plates [in russian and latin]. Chun, W. Y. and K. Z. Kuang. 1962. De genere Cathaya Chun et Kuang. Acta Bot. Sinica 10(3):245247, plates 13. [in latin and Chinese]. 22 Arnoldia 66\/3 Cathaya Comes to the Arnold Arboretum o Peter Del Tredici n october 21, 1998, like a bolt out of the blue, the Arnold Arboretum received an unsolicited packet of nearly 600 seeds of the extremely rare Chinese conifer Cathaya argyrophylla from the Fairy lake Botanical Garden in the city of Shenzhen, Guangdong Province, China. We were excited about getting these seeds for two reasons: first, Cathaya argyrophylla is an endangered species endemic to China, with only limited distribution outside that country, and second, we had received seeds three years earlier, in 1995, but to our great disappointment they had failed to germinate. When the Cathaya seeds arrived at the Arboretum they had no markings other than the name of the plant and the return address. It was all rather mysterious, and it wasn't until nearly three years later, during a chance encounter at the new York Botanical Garden, that I met Dr. li Yong who told me that the seeds had been collected from wild trees growing in Zi Yuan County in Hunan Province, and that he had sent them to the Arnold Arboretum. needless to say, I thanked him profusely for the wonderful gift. The day after the seeds arrived at the Arboretum, Jack Alexander and I counted and divided them up into various lots to test their germination following various periods of moist stratification in the refrigerator. Because we could find no written information about the dormancy requirements of the seeds, and because the species is native to a warm temperatesubtropical area, we made the assumption that the seeds probably required minimal chilling. Table 1 lays out the parameters and results of the seed germination experiment we set up in the Dana Greenhouses. Number of seeds 200 100 100 159 Percent germination (Number of seedlings) 6 (12) 21 (21) 31 (31) 74 (118) Number of days to first seed germination 170 24 29 18 Number of days of chilling 0 57 70 112 Table 1. Germination of seeds of Cathaya argyrophylla which were sown or moist stratified on october 22, 1998. Despite our best guess, the seeds which received four months of cold stratification germinated much faster and in much higher percentages than seeds which received less than seventy days of chilling. So much for a propagator's intuition. By the time the experiment ended in July 1999, we had potted up a total of 182 seedlings, which made for an overall germination rate of 32.6%. Cathay Silver Fir 23 This ten-year-old Cathaya argyrophylla was transplanted to the Arboretum grounds in spring 2008. From the botanical perspective, this species is intriguing because it occupies an intermediate position within Pinaceae, sharing certain morphological similarities with true pines (Pinus), Douglas firs (Pseudotsuga), and spruces (Picea). on April 5, 2000, the Arboretum distributed 79 seedlings to various botanical gardens throughout the United States, keeping about a dozen plants for ourselves. Several of our plants grew well; by spring of 2006, after eight growing season, the three biggest plants were 1.2, 0.8, and 0.7 meters (4, 2.6, and 2.3 feet) tall. Five of the biggest seedlings were moved from the shade house to the nursery in June of 2006, but three of them failed to survive the transplanting. only one plant was still alive by April 2008, when it was planted out on the grounds. our fingers are crossed that it will survive its first winter out on the grounds. As for the seedlings that we distributed back in 2000, the Mendocino Botanical Garden and the University of California Botanical Garden in Berkeley have both reported having plants that are still alive. Peter Del Tredici is a Senior research Scientist at the Arnold Arboretum. PETEr DEl TrEDICI 24 Arnoldia 66\/3 An Excerpt From: Three Conifers South of the Yangtze ur final goal was to reach the Jinfu Shan, the mountainous home of the extraordinary conifer Cathaya argyrophylla . . . After a good night's rest in a fairly decent hotel in nanchuan, we eagerly headed to the jeeps for the drive up into the Jinfu Shan. To our surprise, blocking the gate to the hotel were at least a dozen people arguing with Dr. Yin and Professor Zhong. Apparently several of them were determined to keep us from visiting the Cathaya. There was a representative from the local police, the local tourist bureau, the forestry department, the public security bureau, the Chinese army, the mayor's office, and who knows what else. All were yelling and throwing their arms up in the air. Finally they agreed that we could go see the trees but stated emphatically that we would not be allowed to touch or photograph them. At this point the argument was on the verge of getting seriously out o f c o n t r o l . D r. Yin then made a phone call to the governor, who told the troublemakers that we could indeed visit and photograph the valuable resource Yinshan, the Chinese name for Cathaya argyrophylla, as we were important scientists from England and America. Two and a half The expedition's reward: seeing the silver-backed foliage of Cathaya argyrohours later our phylla in person. jeeps, with an escort of six Chinese to keep us under control, were climbing up steep, mist-covered mountains. We stopped at about 1700 meters (5600 feet) elevation in an area of dense bamboo . . . We then hiked in a light rain for about 20 minutes, slightly uphill, to a large limestone outcrop about 15 meters (50 feet) high and wide. our Chinese escorts pointed to the top of the outcrop and said, \"There they are.\" Through the mist we could barely make out several conifers growing on the top. As we stood there wondering if they would let us climb up to view them closer, we noticed that someone had already rendered that nearly impossible. Everywhere that it might have been possible to climb, the limestone outcrop had been o William McNamara WIllIAM MCnAMArA Cathay Silver Fir 25 (continued from page 21) Fu, l., n. li, and r. r.Mill. 1999 Cathaya. In: Flora of China vol. 4, p. 37. Science Press, Beijing\/ Missouri Botanical Garden Press, St. louis. Fu, l. K. et al. 1992. China Plant Red Data Book--Rare and Endangered Plants. vol. 1. Science Press, Beijing. Haas, W. J. 1988. Transplanting Botany to China: The Cross-Cultural Experience of Chen Huanyong. Arnoldia 48(2):925. Arnold Arboretum, Jamaica Plain, MA. Krussman, G. 1985. Cathaya. In: Manual of Cultivated Conifers. pp. 17, 6263. Timber Press, Portland. li, W. and X. Zhao. 1989. Cathay Silver Fir in Huaping and Jinfo Mountains nature reserves. In: China's Nature Reserves, pp. 128131. Foreign languages Press, Beijing. (brief mentions also of Cathay Silver Fir on pp. 7, 12, 15, 16, 113. Also in Appendix pp. 167, 168, 172, 174, 182). rushforth, K. 1987. Cathaya. In: Conifers. p. 93. Christopher Helm, london. Spongberg, S. A. 1995. Cathaya comes to the Arnold Arboretum, Arnoldia 55(3): Fall news 12. Tang, X. 1987. The Secret Cathay Silver Fir. In: Living Treasures. An Odyssey Through China's Extraordinary Nature Reserves. pp. 8394, illustration p. 196. Bantam Books, new York\/ new World Press, Beijing. van Gelderen, D. M. and J. r. P. van Hoey Smith. 1996. Cathaya. In: Conifers: The Illustrated Encyclopedia vol. 1 AK p. 8, photo p. 90. Timber Press, Portland. Wu, Z., and P. H. raven with G. Zhu (eds.) 2001. Cathaya argyrophylla. In: Flora of China, Illustrations, vol. 4. p.41. Science Press, Beijing\/ Missouri Botanical Garden Press, St. louis. Yu, D. (ed.) 1983. The Botanical Gardens of China. Science Press, Beijing. (photos of Cathaya argyrophylla on pp. 190, 248). Zhang, C. (ed.) 1990. Natural Resources: Vegetation and Plants in the People's Republic of China (Profile China Series). pp. 6377. lotus Publishing House, Hong Kong, and Carlingford, Australia. (Cathaya argyrophylla mentioned pp. 64, 72 with photos both pages). Zhao, J. et al. 1990. The Natural History of China. Collins, london. (Cathay Silver Fir mentioned pp. 52, 193, 199, 209210). Christopher B. Callaghan is a plant researcher and Curator of living Collections at the Australian Bicentennial Arboretum. altered to prevent that possibility. Cracks that might have been footholds had been filled in with concrete, rough areas that might have served as grips were smashed smooth, and in areas of easy accessibility, barriers of rock and concrete had been installed. Someone was undoubtedly determined to keep people away from the Cathaya. As we looked around, clearly frustrated and not trying very hard to disguise it, the Chinese surprised us all by picking up a small fallen tree and leaning it against the outcrop. They then found another similar log and together with the other, they created a makeshift ladder. Several minutes later, after pushing and pulling each other up onto the top of the outcrop, we were standing in a grove of Cathaya. our hosts further surprised us by telling us that it was all right to climb the trees and to take an herbarium specimen. The dozen or so trees averaged about 10 meters (33 feet) in height and superficially resembled short-needled pines... After a good half hour of climbing, examining, and photographing the trees, we slowly made our way back down the outcrop. The rain intensified as we walked back to the road. While getting into the jeeps, our escorts told me that I was the first American to see Cathaya argyrophylla in the wild. Though very suspect of that statement, and rather cold and wet, I was nonetheless very happy to have seen, photographed, and even climbed the Cathaya. William Mcnamara is Executive Director at Quarryhill Botanical Gardens in Glen Ellen, California. Full article at: http:\/\/www. quarryhillbg.org\/page16.html "},{"has_event_date":0,"type":"arnoldia","title":"The Li Jiawan Grand Ginkgo King","article_sequence":4,"start_page":26,"end_page":30,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25453","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14eb36f.jpg","volume":66,"issue_number":3,"year":2009,"series":null,"season":null,"authors":"Xiang, Zhun; Xiang, Yinghai; Xiang, Bixia; Del Tredici, Peter","article_content":"The Li Jiawan Grand Ginkgo King Zhun Xiang, Yinghai Xiang, Bixia Xiang, and Peter Del Tredici he largest Ginkgo biloba tree in the world, the Li Jiawan Grand Ginkgo King, is located about a hundred kilometers west of Guiyang, the capital of Guizhou Province, China. The tiny hamlet of Li Jiawan (2639' N and 10725' E) is too small to appear on any maps. Administratively, Li Jiawan is part of Lebang Village, which is part of Huangsi Town in Fuquan County. The Grand Ginkgo King is growing at an altitude of 1,300 meters (4,265 feet) in a narrow valley where it towers over the surrounding bottomland vegetation, which consists mainly of cultivated crops (Figure 1). It is a male tree, about 30 meters (98 feet) tall, with a ground level trunk diameter of 460 centimeters (181 inches) in the eastwest orientation and 580 centimeters (228 inches) in the north south direction. Its circumference at breast height is 15.6 meters (51 feet) and its canopy shades an area of roughly 1,200 square meters (13,000 square feet). The primary \"trunk\" is completely hollow and encloses an area of 10 to 12 square meters (108 to 130 square feet), more than enough for seating a dinner party of ten people. Indeed, during the 1970s, an old man by the name of Pan Shexiang, accompanied by his cattle, lived in this natural tree cave for two years. The inside of the trunk--up to a height of about 5 meters (16 feet)--is charred black from lightning-ignited fires (Figure 2). The outside of the trunk shows no signs of fire, but has a ragged appearance caused by the excessive amount of callus tissue that has formed between the new branches and old trunks. In addition, large hanging chichi (downward growing shoots that look something like stalactites) have developed in response to various wounds and breaks, adding more confusion to the convoluted woody excrescences that cover the trunk. As battered as the outside of the tree appears, however, it maintains a vigorous hold on life, as attested to by the presence of T numerous young shoots sprouting out all over the tree (Figures 2 and 3). Chinese investigators have determined that the Grand Ginkgo King is a \"five-generationsin-one-tree\" complex. In other words, the first generation was a normal seedling which--as a result of repeated sprouting from the base over the course of several millennia--produced four succeeding generations of trunks, each of which has continued the tree's growth and development after the preceding generation was damaged or died (Figure 4). The tree, as we know it today, is the result of at least five generations of stems produced over the course of thousands of years. There are five distinct trunk sectors which are separate at ground level but are partially merged at the height of about a meter (3.3 feet) above the ground, and new branches often sprout from the tissue between trunk sectors. While each trunk section seems to be physiologically independent, the secondary fusion creates the appearance of a single tree (Figures 2 and 3). Age Estimation Extensive field work has shown that the Li Jiawan Grand Ginkgo King is the biggest (in terms of trunk diameter) ginkgo tree in the world, a fact what was recognized by the Guinness Book of World Records in 1998. The question of how old the tree might be is unclear given that its internal tissues--with all their growth rings--are totally gone. What we do know, however, is that ginkgo trees of different ages have very different appearances and growth characteristics, and that different generations of ginkgo trunks typically have different growth rates and different longevities. We have come up with a rough estimate of the Grand Ginkgo King's age based on what we know about the ages of other ancient ginkgo trees in China with a similarly complex developmental history: the first generation stem(s) can typically reach up to 1,200 years of age, the Grand Ginkgo King 27 P. DEL TREDICI. Figure 1. The Li Jiawan Grand Ginkgo King as it appeared in September 2002. second generation stems live for about 1,000 years, the third 800 years, the fourth 600 years, and the fifth about 400 years. According to this highly theoretical formula, the Li Jiawan Grand Ginkgo King has a maximum estimated age of around 4,000 to 4,500 years. Legends and Romance The Grand Ginkgo King has been living for thousands of years without an official record in the history books of the local government. However, there are many folk legends surrounding this tree. Writer Shixian Xu described one of these legends: During the Tang dynasty there was a scholar named Bai who had recently gained a governor's position by winning a national competition. At some point after taking office, Bai had a fight with a treacherous court official who had done a lot of bad things to the ordinary people. Given that bad officials typically protect each other, the scholar Bai was punished for his actions and sent off to an isolated army camp. On the way there, he was severely beaten and eventually died from his wounds. His body was buried at Li Jiawan by the local people, who deeply loved this scholar who tried to help ordinary people. Soon afterwards, a huge tree grew out from the tomb. This tree was considered the avatar of scholar Bai and given the name \"bai guo tree\" (one of the Chinese names for Ginkgo biloba). Another story about the origin of the tree dates from the Ming dynasty and holds that the Li Jiawan Grand Ginkgo tree transformed itself into a scholar and entered a national competition. The tree-scholar won the championship and was appointed to be a high official by the king. When the tree-scholar failed to show up for the position, the king sent two messengers to find him, both of whom were killed when they came back empty handed. The third messenger that the king sent was worried about his own safety since he too could find no trace of the mysterious scholar. During his disturbed sleep one night, he had a dream in which a person appeared calling himself \"Bai.\" At this point 28 Arnoldia 66\/3 DRAWING By yINGHAI XIANG. Figure 2. The Li Jiawan Grand Ginkgo King. Grand Ginkgo King 29 P. DEL TREDICI. Figure 3. The multi-generational trunk of the Li Jiawan Grand Ginkgo King. 30 Arnoldia 66\/3 DRAWING By yINGHAI XIANG. there have been no new sprouts from part 3. Such a loss of normal regenerative function suggests that the Li Jiawan Grand Ginkgo may be losing its vigor. Based on what we have seen of other multigenerational trees, it is predictable that the Li Jiawan Grand Ginkgo will get smaller over time rather than bigger and that in 50 to 100 years or so it will be dead. References Xiang, B., Z. Xiang, and y. Xiang. 2007. Report on wild Ginkgo biloba in Qianzhong altiplano. Guizhou Science 25(4): 4755. Xiang, B., Z. Xiang, and y. Xiang. 2006. Investigation of wild Ginkgo biloba in Wuchuan County of Guizhou, China. Guizhou Science 24(2): 5667. Xiang, y. and B. Xiang. 1997. Primary report on ancient Ginkgo biloba remnant community in Wuchuan county of Guizhou Province. Guizhou Science 15(4): 239244. Xiang, y., X. Lu, and B. Xiang. 1998. Ancient Ginkgo biloba report 2: data of ancient Ginkgo biloba remnant communities in Luping village and Fengle town of Wuchuan county Guizhou Province. Guizhou Science 16(4): 241252. Xiang, y. and Z. Xiang. 1999. Ancient Ginkgo biloba report 3: investigation on ancient Ginkgo biloba remnant population in Guiyang. Guizhou Science 17(3): 221230. Xiang, Z. and y. Xiang. 2001. Ancient Ginkgo biloba report 4: investigation of ancient Ginkgo biloba remnant population from Changming to Jingyang along 320 national highway in Guizhou province. Guizhou Science 19(1): 4858. Xiang, y., B. Xiang, M. Zhao, and Z. Wang. 2000. A report on the natural forest with Ginkgo biloba population in West Tianmu Mountains, Zhejiang Province. Guizhou Science 18(1-2): 7792. Xiang, Z., Z. Zhang, and y. Xiang. 2001. Investigation of natural Ginkgo biloba population on the Golden Buddha Mountains of Nanchuan, Chongqing. Guizhou Science 19(2): 3752. Xiang, Z., Chenglong Tu, and yinghai Xiang. 2003. A report on Ginkgo resources in Panxian county, Guizhou province. Guizhou Science 21(1-2): 159174. Zhun Xiang is Research Assistant at Guizhou Academy of Science in Guizhou and Graduate Student at South China Agriculture University in Guangzhou. yinghai Xiang is Professor of Ecology at Guizhou Academy of Science in Guizhou. Bixia Xiang is Assisstant Professor of Genetics at the University of Miami. Peter Del Tredici is a Senior Research Scientist at the Arnold Arboretum. Figure 4. A cross-section of the Li Jiawan Grand Ginkgo King at ground level: Part 1 has two trunks: A, 30 meters (98 feet) tall, 110 centimeters (43 inches) diameter; B, 20 meters (66 feet) tall, 90 centimeters (35 inches) diameter; Part 2 has produced trunk C with a height of 28 meters (92 feet) and diameter of 80 centimeters (31 inches); Part 3 has trunk D of height of 28 meters (92 feet) and diameter of 60 centimeters (24 inches). The smallest and youngest trunks, Parts 4 and 5, have produced many small, weak stems, only a few meters tall, which seem to have lost their capacity to grow into upright trunks. the messenger woke up and saw an official's hat hanging on the top of a nearby ginkgo tree and immediately understood that the scholar and the tree were one and the same. This story--that the ginkgo tree had changed to a spirit--is an astonishing, age-old story, and there are lots of \"big tree changed to spirit\" stories in the south of China. Luckily, people usually worship such \"spirit trees\" and don't dare to damage them. Many of these trees grow in temple courtyards or on sacred mountains and are preserved out of respect for the spirits that inhabit them but, unfortunately, this kind of conservation is not good enough to protect trees in the modern world. What the Future Holds The Li Jiawan Grand Ginkgo King was seriously damaged and its overall appearance dramatically changed by a storm in July, 1991, in which the biggest trunk on part 2 was broken off (Figure 4). The stem was pruned off below the break, but the resulting scar still looks fresh with no sign of callus growth to cover it over. It is also worth noting that for eighteen years "},{"has_event_date":0,"type":"arnoldia","title":"An African Fir Grows in Boston","article_sequence":5,"start_page":31,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25450","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14eab6d.jpg","volume":66,"issue_number":3,"year":2009,"series":null,"season":null,"authors":"Port, Kyle","article_content":"An African Fir Grows in Boston Kyle Port T he Arnold Arboretum's Conifer Collection offers visitors an opportunity to explore gymnosperms collected from around the world. While Eastern Asian, European, and North American species dominate the collection, a solitary Moroccan fir, Abies pinsapo var. marocana, stands as an exceptional North African taxon. Grown from seed collected by former Arboretum plant propagator Rob Nicholson on Mt. Tisouka near Chefchaouen, Morocco, in 1982, specimen 1435-82-A has thrived undamaged in the landscape for 15 years. It is one of two plants of accession 1435-82 that were moved from the Arboretum's Dana Greenhouse to the grounds on September 21, 1993. One plant did not survive transplantation and was noted as dead in the spring of 1994. The lone survivor, which was approximately 4.3 feet (1.3 meters) at the time of transplant, is now a stunning exemplar at 28 feet (8.5 meters) tall with a DBH (diameter at breast height) of 12.6 inches (32 centimeters). Conical in youth, the tidy habit of this specimen has opened slightly over the years to reveal smooth gray bark. Radially arranged needles persist for 11 to 13 years, giving even older branches an armored appearance. The dark green needles are streaked with 7 to 11 silvery stomatic lines on the upper surface; the lower surface is marked with two pronounced stomatic bands on either side of the midrib. Unlike the characteristically soft-to-the-touch foliage of most Abies, the needles of Abies pinsapo var. marocana have sharply pointed apices, making the foliage far less friendly to fingers. The upright cylindrical cones typical of the species have not yet been observed on this specimen but can be expected soon; sexual maturity for Moroccan fir is typically reached when the trees are between 25 and 35 years old. Described by French botanist Louis Charles Trabut in 1906 as A. marocana, the Moroccan fir is confined to the Rif Mountain Range of Morocco, growing at altitudes between 4,600 and 6,900 feet (1,400 to 2,100 meters). The calcareous soil of this region supports associated taxa, and notes from the Arnold Arboretum's collecting trip detail an open fir forest containing Cedrus atlantica, Acer (A. opulus ssp. hispanicum, A. campestre, A. monspessulanum), and Paeonia (P. coriaceae var. maroccana). Rare in cultivation, the International Union for Conservation of Nature and Natural Resources considers Abies pinsapo var. marocana to be a \"near threatened\" species, an indicator that it could become threatened in the wild in the near future. Human activities (logging, expansion of cultivated areas, population growth) and climate change may further restrict the range of this taxon. However, preservation efforts are ongoing and the establishment of the Talassemtane National Park, which contains the only remaining Moroccan fir forest, was celebrated by conservation organizations in 2004. Related species: Abies pinsapo `Glauca', blue Spanish fir, is also represented in the Arboretum's collection (accession 192-42-A, obtained from W.B. Clarke and Company, San Jose, California). Planted in the fall of 1954, this blue-hued cultivar is topped with dozens of cones this year. Separated from the Moroccan fir by the Straits of Gibraltar, the Spanish Fir (Abies pinsapo var. pinsapo) is endemic to the Sierra de Ronda in Southern Spain. Kyle Port is Manager of Plant Records at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23411","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd25e896e.jpg","title":"2009-66-3","volume":66,"issue_number":3,"year":2009,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Dysfunctional Root Systems and Brief Landscape Lives: Stem Girdling Roots and the Browning of Our Landscapes","article_sequence":1,"start_page":2,"end_page":10,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25446","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14ea326.jpg","volume":66,"issue_number":2,"year":2008,"series":null,"season":null,"authors":"Johnson, Gary","article_content":"Dysfunctional Root Systems and Brief Landscape Lives: Stem Girdling Roots and the Browning of Our Landscapes Gary Johnson Consider this comparison of potential life spans for trees (Burns and Honkola 1990; USDA 1998) Quercus macrocarpa (bur oak), in upland site Acer saccharinum (silver maple), in riparian site Acer negundo (boxelder), in lowland site Pinus banksiana (Jack pine), in field site Betula papyrifera (paper birch), in northern lowland forest Tree planted in urban core street site 250+ years 125+ years 100+ years 80+ years 65+ years less than 10 years hat's a sobering thought--a tree with a normal life span of 65 to 250 years may live less than 10 years when planted in any American city's downtown landscape. Admittedly, that figure represents tree placement in the worst of our urban landscape sites: sidewalk cut-outs. These inhospitable planting sites are also known as tree coffins, tree burial mounds, or urban tree disposal units to frustrated urban foresters. When the mortality rate of downtown trees is compared to tree losses from Dutch elm disease, oak wilt, sudden oak death, and gypsy moth, it doesn't take too long to realize that there's an epidemic of urban tree loss going on and it's largely under the radar (Figure 1). Another oft-quoted number is that the average urban residential tree lives for 30 to 35 years (Moll 1989). That life span is three times as long as a sidewalk tree, yet only half as long as a paper birch in its natural environment. Growing conditions in residential landscapes may not be quite as bad as sidewalk sites, but there are many natural and unnatural pressures on the trees that lead to briefer landscape lives. Residential landscape soils can be as stressful as downtown sites: poorly drained, outrageously alkaline, subjected to blends of every pesticide T known to modern society, and compacted to such a degree that lawns may seem like nothing more than green concrete. With few exceptions (perhaps tornadoes and a few diseases), there are no \"angels of death\" that descend and quickly kill trees in landscapes. More commonly, a multitude of predisposing stresses that occur in our highly altered urban landscapes combine to weaken trees over the years. Often, inciting events such as floods or hailstorms and\/or contributing agents such as target cankers or wood boring insects complete the job for the majority of tree losses. Meanwhile, plant health care professionals attempt to determine the true causes of decline and death, and often the diagnoses are incomplete or incorrect because of the multiple offenders involved with the problem. Predisposing Factors and Tree Decline When trees are chronically stressed (long-term drought, repeated defoliation, etc.), their normal reserves of chemical energy--primarily as complex carbohydrates--are slowly depleted. Each year as stressed trees come out of dormancy, they emerge in a weakened state due to this energy depletion and find it increas- Dysfunctional Root Systems 3 PHOTO By GARy JOHNSON Figure 1. Trees in urban sidewalk sites are subjected to very unhealthy environments and live less than 10 years on average. ingly difficult to releaf, grow, and deal with the harsh realities of urban landscapes on a normal basis. It takes a tremendous amount of chemical energy to push out new leaves and shoots, recover from accidental wounds on the stems, or produce flowers and fruit. As the tree's energy reserves continue to decline--and thereby affect the tree's ability to capture and store new energy through photosynthesis--the entire system is affected and the decline spiral to premature death begins. So decline in a sense refers to the tree's ability to deal with life's normal stresses. A tree in decline may die suddenly because of an event such as a cold winter with no snow cover, a short-term summer drought, or a defoliation from insects or hail. The other trees in the landscape tolerate the damage and survive, but the predisposed trees--those in decline--are unable to recover from the damage. Dysfunctional Root Systems as Predisposing Agents Despite the fact that roots are seldom seen, dysfunctional root systems are too often the predisposing agents connected to tree health decline, and ultimately the reason why many urban landscape trees experience such brief lives. If the root system--approximately 50% of a tree's biomass--is not operating normally, the entire system will be abnormal. Abnormal is not always harmful, as seen in bonsai plants and trees growing on slopes. In bonsai plants, a restricted root system causes compacted growth in the rest of the plant system, but the system itself may be healthy and completely functional under most circumstances. In the case of a tree growing on a slope, the tree is anchored with a skewed and asymmetrical root system, but its overall health is not compromised even though the root system could certainly be considered abnormal. 4 Arnoldia 66\/2 PHOTO By DAVE HANSON which create a root system so dysfunctional that it can end up killing the entire tree. Stem Girdling Roots as Predisposing Agents Stem girdling roots are those roots that grow either partially or completely against the tree's stem and compress (girdle) the stem tissues (Figure 2). Xylem and phloem tissues in the stem become much narrower at the point of compression, impeding normal water movement and sap flow (Figure 3). This restriction affects energy reserves by directly Figure 2. With part of a stem girdling root removed, the compression to the and indirectly affecting photosyntree's trunk is evident. thesis. Trees become stressed and But abnormal root systems that do affect more vulnerable to secondary problems. For the overall health or stability of the tree are this reason, SGRs are considered to be primary considered dysfunctional. For example, when a predisposing agents in landscape tree decline container-grown tree with a severely pot-bound and death. root system is planted, its rhizosphere does not Some of the first symptoms of SGR-impacted occupy a large enough area to capture sufficient tree health include leaf scorch or leaf wilting water and nutrients needed to support a normal on a tree when no other plants in the area are sized tree without supplemental help. Dysfuncshowing the same symptoms. There may be tional root systems are also common on newly adequate moisture in the soil, but the tree's transplanted bare-root and balled-and-burlapped ability to move water throughout the system is plants; these plants often lose 75% or more of thwarted by the areas of compression, i.e. the their root systems during the harvest operation, greatly reduced diameter of vessel elements. resulting in transplant shock which may go on Soon, this water stress evolves into early leaf for several years until the root system regrows. coloration and leaf drop in the summer, late And then there are stem girdling roots (SGRs), leaf-out in the spring, and chlorosis or other G. W. HUDLER, CORNELL UNIVERSITy Figure 3. Transverse views of normal Norway maple stem wood showing a healthy growth pattern (left), and malformed stem wood compressed by a stem girdling root (right). Water and nutrient transport in trees is negatively affected when tissue is malformed by compression. V = vessel element, R = ray, F = fiber tracheid. Both views are at the same scale. Dysfunctional Root Systems 5 PHOTOS By GARy JOHNSON nutrient deficiency symptoms. If the stem compression becomes more severe, affecting 50% or more of the stem circumference, so do the symptoms. Trees will tend to suffer more damage during the winter seasons, in particular true frost cracks, cambial death, and dieback. In the latter stages of decline due to SGRs, trees usually suffer from severe stunting (very small leaves, annual twig growth of 1 to 2 inches or less) and significant defensive dieback. With Figure 4. The middle littleleaf linden was in the last stages of decline from stem so little vascular capacity left, girdling roots at the time of this photograph. One year later it was dead. affected trees may succumb completely from even a short-term summer drought (Figure 4). Though often a slow-acting cause of death, SGRs can also cause tree death that is a bit more sudden and dramatic. The compressed areas of tree stems are structurally weak points and far too often are the points of failure during windstorms (Figure 5). For example, in severe windstorms that occurred in Minnesota in 1998, 73% of the lindens (Tilia spp.) that were lost in urban Figure 5. Stem compression from SGRs located 4 or more inches below ground landscapes failed at compression was the most common cause of urban tree failure in windstorms in Minnesota points from SGRs, and most broke from 1995 to 2005. several inches below ground. This these SGRs were well below ground (from 4 to is a different type of predisposition but equally 14 inches)--out of sight, out of mind (Figure 6). damaging to a tree's ability to grow, survive, and In landscape surveys conducted by the Uniadd to the quality of life. versity of Minnesota Department of Forest More (Soil) is Not Always Better Resources (1997 to 2004), five species of trees were investigated in three different communiEarly SGR studies conducted by the University of ties. All trees were growing in public spaces: Minnesota were in response to unexplained tree boulevards, schools, government centers, parks. decline in urban areas. From 1994 through 1996, Species surveyed included hackberry (Celtis 220 declining and dying trees were diagnosed. In occidentalis), littleleaf linden (Tilia cordata), 81% of the cases, stem girdling roots were the sugar maple (Acer saccharum), `Shademaster' only causal agents isolated. This figure closely honey locust (Gleditsia triacanthos `Shademasparalleled data collected from a national survey of ter'), and green ash (Fraxinus pennsylvanica). tree care professionals (Johnson and Hauer 2000). Trees were randomly selected, evaluated for More specifically, these trees had been planted health and condition, and then examined for in the previous 12 to 20 years and had signifidepth of soil over the main order roots and the cant stem compression (greater than 50% of the presence of stem encircling roots (potentially stem circumference) from SGRs. In all cases, 6 Arnoldia 66\/2 conflicting roots within 6 inches of the stem) or stem girdling roots. The results were a bit depressing. Only 4% of the lindens, 8% of the ash, 10% of the maples, 15% of the honey locust, and 40% of the hackberries had their stems completely above ground. The rest of the sampled trees had from 1 to 12 inches of soil over the first main order roots and against the stems. PHOTOS By GARy JOHNSON Non-destructive root collar examinations were performed on a total of 1,380 trees. The intent of these examinations was to determine the frequency of SERs (stem encircling roots-- those potentially conflicting roots within six inches of the stem) and SGRs associated with different depths of soil (up to 12 inches) over the first main order roots. The excavations demonstrated that the deeper tree stems were buried in the soil or mulch, the more likely it was for them to have multiple layers of stem encircling and stem girdling roots. The increased presence of these problem roots showed up in trees beginning with as little as one inch of excess soil against the stem. In a nutshell, the more soil or pre-soil (organic mulches that will break down) that is piled over the root systems and against the stems, the more likely it is that trees will decline or fail due to multiple conflicts with SGRs (Figure 7). Figure 6. This SGR, located approximately 4 inches below ground, runs tangential to the tree trunk and is compressing 30% of the stem circumference. Figure 7. As shown on this littleleaf linden, more layers of SGRs develop as the stem is buried deeper. Greater than 40% of the stem circumference of this tree was compressed by several layers of SGRs. How SGRs Form Observations from the 1,380 root collar examinations conducted during the species surveys and a separate nine-year planting depth study have led to the conclusion that stem girdling roots form in one of two ways: first, new roots regenerating from deeply buried main order roots, and second, from stem adventitious roots. When main order roots are buried too deeply, new woody roots that originate from them or any part of the buried root system tend to grow closer to the surface. It is speculated that this action is in response to a more desirable soil oxygen and moisture balance. As the roots reach the soil surface, an unpredictable percentage of them grow tangential to the tree stem or in some cases encircle the stem. For the next number of years (12 to 20, from our observa- Dysfunctional Root Systems 7 tions), the roots and stems expand in diameter, resulting in the ultimate confrontation between roots and stems. Stem adventitious roots are also sources of SGRs. When a buried stem begins forming adventitious roots, many or most of those roots grow away from the stem in a radial fashion. As with new roots growing from main order roots, an unpredictable percentage of these adventitious roots do not grow radially but instead grow tangential to the stem or encircling the stem. The interface area between soil and stem appears to be a highly desirable area for stem root growth, perhaps because it provides an ideal balance of soil oxygen and moisture and is also the path of least resistance for root proliferation. The exact reasons for these root growth responses are still speculative, but it is clear that when tree stems are buried by a media that supports root growth, SGRs are highly likely to occur. It's worth noting that stem girdling roots are a problem primarily with younger trees. As trees mature, their growth slows down dramatically, including the growth of trunk diameter and encircling roots. Because of this reduced growth--and the fact that there is often a relatively thick outer bark--stems of mature trees that then become buried by soil or organic matter are much less likely to develop stem girdling root problems. SGRs can still develop, but if they do they are less likely to result in the decline and death of the tree. Trees are planted in a new landscape before final grading is completed. There are so many different ways that stems can be buried--accidentally or with good intentions--that it is difficult to pinpoint the main source of the problem. One seemingly common cause is the act of burying trees rather than planting trees. Unfortunately, too many people still have the notion that trees are like fenceposts and need to be buried deep for stability. Not so. In 2002, we conducted a planting depth study in collaboration with a large wholesale nursery. Bare-root birch (Betula spp.), ash (Fraxinus spp.), and crabapple (Malus) were potted up in number-ten containers at four different depths: 0, 2, 4, or 6 inches of soil over the first main order roots. On a weekly basis, each of the 240 trees was inspected for lean or windthrow from the containers. At the end of the four month study, all trees were well-rooted in the containers and the results of the study showed that all trees, regardless of depth, leaned at the same frequency and to the same degree. Planting tree stems deeper had absolutely no positive effect on tree stability. If newly planted trees are unstable, they may need temporary support from a guying or staking system, not entombment. How to Cause Stem Girdling Roots If you want to cause the formation of SGRs, bury the tree stem with a medium that supports root growth. Here are some common ways SGRs occur: Excess soil is piled over the first main order roots during the growing and harvesting of balled-and-burlapped trees. Excess growing medium buries stems when container-grown trees are up-potted. Decayable organic mulch is piled high around tree stems in nurseries and landscape sites. Soil is piled against tree stems during construction regrading in landscapes. Nine Years of Burial In 2000, a long-term planting depth study was installed at the University of Minnesota's Urban Forestry and Horticulture Institute's research fields. Three hundred and sixty trees equally represented by two species (sugar maple [Acer saccharum] and littleleaf linden [Tilia cordata]) were planted at three depths: 0, 5, or 10 inches of soil over the first main order roots. All trees were planted in a complete, randomized block design in a .75 acre plot as unbranched, 2 to 3 feet tall liners. At three year intervals, onethird of the trees were harvested and had their root systems excavated with a supersonic air tool. Each year, mortality rates, growth rates (stem caliper), number of suckers produced, and percentage of dieback was recorded. In 2009, the final third of the original experiment will be harvested, but some interesting trends and 8 Arnoldia 66\/2 PHOTO By GARy JOHNSON PHOTO By NANCy ROSE Figure 8. Bury the stem of littleleaf linden just 5 inches deep and a profusion of suckers will develop. These suckers eventually become SGSs (stem girdling suckers) as they grow in caliper and compress the tree's stem. significant data have already been revealed from the first two harvests, including: Planting sugar maples 5 to 10 inches too deep is an effective way to kill them. The mortality rates for the 0, 5, and 10 inch depths as of 2006 were 30, 40, and 65%, respectively. There was a significant positive relationship between placing 5 to 10 inches of soil against the stems and the frequency of SGRs on Tilia cordata in both the 2003 and 2006 harvests. Acer saccharum showed a trend in the same direction. Tilia cordata with stems buried in 5 inches of soil will produce masses of stem suckers, making the tree look more like a shrub. Sucker formation on Tilia cordata doesn't just ruin the tree's appearance, it can also cause premature failure. Stem girdling suckers (SGSs) are suckers that form prolifically and, when they enlarge in diameter, can girdle the stem vertically and horizontally (Figure 8). How Often do Trees Die from SGRs? This question is likely unanswerable. When trees suddenly fail and die during a windstorm, diagnosing the problem below ground is not often considered. Weather alone is often blamed for the deaths, and the trees are hastily removed and replaced. Research we conducted from 1995 through 2005 on tree failure in windstorms exposed a Dysfunctional Root Systems 9 broader picture of the effects SGRs have on landscape trees. During this period over 1,500 \"tree autopsies\" were conducted on trees that had failed during wind-loading events in Minnesota. These trees were not those from the centers of severe wind-loading events such as straight-line winds or tornados. Rather, they were victims of thunderstorms or those at the edges of severe wind events. From that data, the destruction and economic losses from premature tree failures due to SGRs were determined, and it was startling. The most common tree size category for boulevard tree failures was the 6 to 10 inch DBH (diameter at breast height, 4.5 feet above ground) range. Of those trees, 50% snapped off at compression points from SGRs at a depth of 4 or more inches below ground. The Achilles' heel was a compression root that couldn't even be seen because the stem was buried so deeply. The data also indicated that littleleaf lindens (Tilia cordata) were grossly affected by SGRs. Littleleaf linden ranked as the third most common species for total failure (the tree went down completely) during those years, and 73% of those trees snapped off at below-ground SGRs, almost the exact percentage of littleleaf linden that failed during the previously mentioned 1998 storms. After 11 years of data collection, the presence of SGRs and, more specifically, stem compression from SGRs that amounted to 50% or more of the stem circumference, emerged as the number one reason why urban trees failed in windstorms. What to Do, What to Do? Prevention is the easiest and most effective way to eliminate the SGR problem in landscapes. Whether you are an urban forester, commercial landscaper, or home gardener, follow these steps to prevent or manage stem girdling roots: Don't plant container or balled-and-burlapped trees that are already buried too deeply. Assume there is too much soil over the first main order roots and remove that excess soil before planting a newly purchased tree (Figure 9). Plant trees, don't bury them. If stems aren't buried, it's not likely that SGRs will become a problem. They can still occur on correctly planted trees, but much less frequently than on buried trees. PHOTO By GARy JOHNSON Figure 9. Most containerized trees will have 2 to 6 inches of excess soil over the first main order roots and against the stem. Use a pruning saw to remove this excess soil before planting. Of 500 trees subjected to this treatment at the University of Minnesota's research nursery, there has been a 0.7% mortality rate in 2.5 years. 10 Arnoldia 66\/2 Don't pile mulch against stems. Organic mulch is basically presoil. Piling on mulch will result in a buried stem and a wonderful environment for SGRs to develop. When suspicious, investigate. Root collar exams are not all that difficult to perform (Figure 10). If you have a trowel and a wet-dry vacuum, you can perform a non-destructive root collar exam. If you find offending roots during the exam, remove them. Also, Figure 10. The fastest and most non-destructive method for conducting a root collar exam remove all that extra is with a supersonic air tool that blows the soil away without harming the roots. This root soil. If you do nothing, collar exam was accomplished in approximately fifteen minutes. it will only get worse. References: If greater than 50% of the stem's circumference is severely compressed, it is probBurns, Russell M., and Barbara H. Honkala, tech. coords. 1990. Silvics of North America: 2. Hardwoods. ably best and safest to remove the tree and Agriculture Handbook 654. U.S. Department of start over. Agriculture, Forest Service, Washington, DC. Treatments for affected trees are uncertain. If SERs (stem encircling roots) can be removed before compression begins, that's an excellent and effective treatment. If the SERs have become SGRs and if, during the course of removing SGRs, the stem is wounded, the long-term potential for recovery is uncertain. The study of stem girdling roots is a relatively young science and long-term data on treatment options and efficacy are not there. If 50% or more of the tree's trunk is severely compressed by the SGRs, and if the symptoms included dieback and severe stunt, the tree is probably beyond salvation. If that same tree is ten feet from a house or utility line, then the risk of leaving the tree is unacceptable. Buy a new tree. Remove the excess soil over the root system. Plant it with the trunk fully exposed. Mulch the roots, not the trunk. These steps will put your new tree well on the way to a long, healthy life. Johnson, Gary and Dennis Fallon. 2007. Stem Girdling Roots: The Underground Epidemic Killing our Trees. www.forestry.umn.edu\/extension\/. Under \"Hot Topics\" on the home page, access \"Stem girdling roots.\" Johnson, Gary and Richard Hauer. 2000. A Practitioner's Guide to Stem Girdling Root of Trees: Impacts on Trees, Symptomology, and Prevention. www.forestry.umn.edu\/extension\/. Access the \"Urban Forestry\" folder, then \"Stem Girdling Roots\" under \"Tree Care.\" Moll, Gary. 1989. The state of our urban forest. American Forests. 95(11\/12):6164. Summary of Storm Damage to Urban Trees in Minnesota: 19952005. www.forestry.umn.edu\/extension\/. Access the \"Research\" folder, then \"Storm Damage 19952005.\" USDA Forest Service. 1998. Urban Forest Health: Identifying Issues and Needs within the Northeastern Area. Gary Johnson is a professor in the Department of Forest Resources at the University of Minnesota. PHOTO By DAVE HANSON "},{"has_event_date":0,"type":"arnoldia","title":"Wake Up and Smell the Ginkgos","article_sequence":2,"start_page":11,"end_page":21,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25448","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14ea76f.jpg","volume":66,"issue_number":2,"year":2008,"series":null,"season":null,"authors":"Del Tredici, Peter","article_content":"Wake Up and Smell the Ginkgos ALL PHOTOS By THE AUTHOR EXCEPT AS INDICATED Peter Del Tredici G inkgo biloba is one tree that most Americans--even those with little knowledge of botany--can recognize. There are two reasons for this: first, its fan-shaped leaves are highly distinctive and impossible to confuse with any other tree; and second, it is widely cultivated as a street tree in many urban areas throughout much of the United States. Because of its environmental adaptability, its resistance to pests and diseases, and its general tolerance of inhospitable growing conditions, ginkgo is experiencing a spike in popularity as evidenced by the long rows of them that are showing up in commercial and municipal landscape projects across the country. In this regard, Americans are following the pattern set in Japan where ginkgo accounts for 11.5% of all the street trees growing in that country--more than any other single species (Handa et al. 1997). As well as gaining in popularity, ginkgo has also been experiencing a surge in attention from the scientific community, particularly from the Chinese, for whom the tree has become a national symbol of their botanical heritage. The pur- An allee of ginkgos, about 100 years old, on the campus of Tokyo University. Ginkgo has unmistakable fan-shaped leaves. 12 Arnoldia 66\/2 One of the old ginkgos at Bai Yuan village in Wuchuan County, Guizhou Province. Note the epiphytic ferns growing on its trunk. pose of this article is to acquaint the reader with some of this new information about the plant's unique evolutionary history as well as its ecological role as a plant teetering on the brink of extinction in the wild. Ginkgo's Homeland Questions about the extent of Ginkgo biloba's native range in China--or if native populations even exist at all--have been the subject of debate among botanists for well over a hundred years (Del Tredici et al. 1992, Li et al. 1999). The conflict has only recently been resolved with the help of DNA analyses (Fan et al. 2004, Shen et al. 2005, Wei et al. 2008) which have demonstrated that isolated ginkgo populations located in southwest China, especially around the southern slopes of Jinfo (or Golden Buddha) Mountain in Chongqing Province (2853' N; 10727' E), possess a significantly higher degree of genetic diversity than populations in other parts of the country, indicating native status. The area has a mesic, warm-temperate climate with a mean annual temperature of 16.6C (62F), and a mean annual precipitation of 1,185 millimeters (47 inches), with ginkgos growing mainly between 800 and 1,300 meters (2,625 and 4,265 feet) elevation (Li et al. 1999). In addition to the genetic evidence, there is ecological and cultural evidence which suggests that these populations are wild. Ecological work in Chongqing Province, as well as in adjacent parts of Guizhou Province (Xiang et al. 2006), has identified dozens of small populations of ginkgos which can be considered either to be wild trees growing in the midst of native forest or the remnants of wild populations that have lost their forest context. These ginkgo populations occupy land that usually measures a few hectares at most, and they are surrounded by small villages whose residents practice subsistence agriculture. In areas where livestock has been excluded, spontaneous ginkgo seedlings and saplings are common in the forest understory. In the cultural realm, much of northern Guizhou Province has been settled over the past three hundred years or so by people of Miao descent who, unlike the Chinese of Han descent, have no tradition of consuming ginkgo nuts and therefore have no history of cultivating the tree. While this situation began to change around 1980, cultivation by humans cannot explain the many large ginkgos scattered throughout the area that are not growing near temples. (Ginkgos found near temples are usually human cultivated.) From the ecological\/botanical perspective, wild populations of ginkgo tend to show a number of characteristics which distinguish them from populations of cultivated trees. These differences are summarized in Table 1 (page 14). In addition to the populations around Jinfo Shan, a second area of high genetic diversity for ginkgo occurs in eastern China, in Zhejiang Province, primarily on the slopes of Tian Mu Shan, a sacred mountain with many Buddhist shrines and temples, located about 100 kilo- Ginkgo Sexuality 13 The agricultural terraces in the vicinity of Shan Jiang village in Wuchuan County, Guizhou Province. Over the past several hundred years, these have replaced the mixed coniferbroadleaf evergreendeciduous forest that originally grew there. meters (62 miles) west of the city of Hangzhou. This area, which was the site of one of the first nature reserves in modern China, has long been considered by botanists to be one of ginkgo's wild locations, but only recently--through the work of Wei Gong and her colleagues (2008) at Zhejiang University--has the distinct genetic ancestry of this population been established. In contrast to its very limited distribution as a wild plant in China, ginkgo is widely cultivated throughout the temperate world, across a broad range of moisture, temperature, and topographic gradients. In China, the tree can be cultivated between 25 and 42 N latitude where minimum winter temperatures can reach -32C (-26F) and maximum summer temperatures 42C (108F) (He et al. 1997). Detailed phenological studies in Japan over a fifty year period by Matsumoto and his colleagues (2003) have determined that spring bud break in ginkgo occurs 40 days earlier in the extreme south of the country (30 N latitude) than it does in the far north (43 N latitude) and that autumnal leaf drop happens about 40 days later, making for an effective vegetative growing season range of 170 to 260 days across 13 of latitude. It's no wonder that ginkgo is touted as a paragon of environmental adaptability. Ginkgo Sexuality Ginkgo biloba is a dioecious species, with separate male and female trees occurring at roughly a 1:1 ratio. Ginkgo shows a long juvenile period, typically not reaching sexual maturity until approximately 20 years of age. Male (microsporangeate) and female (ovulate) sex organs are produced on short shoots in the axils of bud scales and leaves. The male catkins emerge before the leaves and fall off immediately after shedding their pollen to the wind. Pollination 14 Arnoldia 66\/2 (Left) A portion of the stand of wild ginkgos at Bai Yuan village in Wuchuan County, Guizhou Province. Note the tall, straight form of the trees indicting that they grew from seed. (Below) Cultivated ginkgos in this orchard show the typical shorter, wide-spreading form. Table 1. The botanical and ecological characteristics of remnant natural ginkgo populations versus cultivated ginkgo populations in China Remnant natural ginkgo populations Sex ratio should be more or less balanced with males at a 1:1 or greater ratio than females. Trees are growing mixed in with numerous other species that are native to the surrounding forest. The growth form of most of the trees is single stemmed with relatively few lower branches (indicative of having grown up from seed). Cultivated ginkgo populations Skewed sex ratio--overwhelmingly female. Few other species growing with ginkgo; if other trees are present, they are typically cultivated for some specific purpose. Low-branched growth form of female trees (indicative of vegetative propagation by cuttings or grafts). typically occurs anywhere from mid-March in areas with mild winters to late May in areas with severe winters. The ovules on female trees are 2 to 3 millimeters (about .1 inch) long at the time of pollination, and are produced mostly in pairs at the ends of long stalks. When the ovule is receptive, it secretes a small droplet of mucilaginous fluid from its apical tip which functions to capture airborne pollen. Retraction of this droplet at the end of the day brings the pollen into the pollen chamber. Once inside the ovule, the pollen grain germinates to release the male gametophyte which attaches itself to the inside wall of the ovule. Here it undergoes a four- to fivemonth-long period of growth and development which is supported by the tissues of the expanding ovule (Friedman and Gifford 1997). Ginkgo Sexuality 15 Ginkgo ovule with pollination drop at tip. Sometime in September or October, depending on the latitude, the development of the male gametophyte culminates with the production of a pair of multiflagellated spermatozoids. In one of nature's most dramatic moments--first described by the Japanese botanist Hirase in 1896--the two microscopic sperm cells must swim, propelled by about one thousand tiny flagella, a full millimeter across a fluid-filled channel to reach the waiting egg cell, where only one can claim the prize. Contrary to what has often been written, fertilization takes place while the ovules are still on the tree and embryo development begins posthaste. The embryo length may range from less than 1 millimeter to 5 millimeters (.04 to .2 inch) at the time of seed drop, which can occur anywhere between September and November, depending on local weather conditions. Once the seeds fall to the ground, the embryo continues to develop until the arrival of cold temperatures (below 10C [50F]), at which point elongation stops. With the onset of warm weather in the spring, the embryo resumes its growth, which culminates in germination in late spring or early summer. gametophyte surrounded by a thick seed coat. The intact seed coat consists of a soft, fleshy outer layer (the sarcotesta), a hard, stony middle layer (the sclerotesta), and a thin, membranous inner layer (the endotesta). The seed, devoid of the famously smelly sarcotesta, is generally referred to as the \"nut\" with dimensions that range from 19 to 30 millimeters by 11 to 14 mm (approximately 1 by .5 inch). Over the past several hundred years, Chinese horticulturists have selected scores of cultivars which produce large and\/or distinctively shaped nuts. Large plantations of these select ginkgo cultivars are common throughout eastern and central China. The putrid odor often associated with ginkgo seeds typically develops only after they have lain on the ground for several days and have begun to rot. The smell is due to the presence of two volatile compounds in the sarcotesta-- butanoic and hexanoic acids (Parliament 1995). The sarcotesta also contains numerous fatty acids and phenolics, one of which, ginkgoic acid, is known to cause allergic contact dermatitis in some people (Kochibe 1997). A Common-Garden Experiment The timing of pollination, fertilization, seed abscission, and germination in ginkgo are strongly affected by the latitude of cultivation as well as by local climate conditions. In the Ginkgo Nuts It is now generally accepted that ginkgo was first cultivated by the Chinese not for religious purposes but rather for its edible seeds, which at maturity are relatively large and nutritious. The seed, as it falls from the tree, consists of an embryo embedded in the tissue of the female Mature ginkgo seeds on a tree at Forest Hills in Boston, Massachusetts. 16 Arnoldia 66\/2 fall of 2002, I undertook a series of commongarden experiments to explore the relationship between the timing of pollination and the timing of germination in ginkgo by cultivating in a common location seeds produced by trees from two different latitudes. One lot consisted of about 500 cleaned seeds from trees that were being cultivated for nut production, which I purchased on September 22, 2002 at Tuo Le Village, Panxian, in southern Guizhou Province, China, (2536' N). For comparative purposes, I collected ginkgo seeds on October 31, 2002 from beneath a number of trees growing at the Forest Hills Cemetery in Boston, Massachusetts (4217' N). When sown in the Arnold Arboretum's heated greenhouse (20C [68F]), the Guizhou seed began germinating on November 12--approximately 58 days after abscission--while the Boston seed did not begin germinating until January 6--some 67 days after abscission. Assuming approximate pollination dates of March 24 for the Guizhou seed and May 17 for the Boston seed, the total time elapsed from pollination to germination under continuously warm greenhouse conditions was 233 days and 234 days respectively, a remarkably confluent result given their different latitudinal origins. A second striking result of the experiment was that only 15% of the uncleaned, outdoorsown Boston seed germinated versus 72% germination for a replicate lot of one hundred seeds washed clean of their smelly sarcotesta. The fact that cleaned ginkgo seeds germinated at statistically significantly higher percentages than those with their sarcotesta intact suggests that animals which consume the seeds--provided they do not crush the thin-shelled nut--might play a role in promoting successful seedling germination (Rothwell and Holt 1997, Del Tredici 2000). The specific mechanism whereby the sarcotesta reduces the germination capacity of ginkgo seed is currently unknown, but the exclusion of light is probably not an explanation given that William Friedman (1986) has shown that female gametophytes with all their seed coats intact are capable of photosynthesis. Ecological Implications The results of my experiment indicate that aspects of ginkgo's sexual reproduction cycle are strongly influenced by temperature (Del Tredici 2007). For seeds left outdoors immediately following seed drop, the timing of their pollination influences the timing of their germination the following spring which, in turn, influences their chances of surviving the following winter. In warm-temperate climates--such as Guizhou Province--ginkgo seeds are shed in late summer or early fall, and the embryo is able to make considerable growth during the mild weather that follows. In cold-temperate climates--such as Massachusetts--seeds are shed much later in the season and the cooler temperatures of mid to late fall delay embryo development until warm weather arrives the following spring. This differential timing of embryo maturation means that seeds produced by trees growing in warm-temperate climates will be ready to germinate during the favorable conditions of Table 2. A comparison of the phenology of the sexual reproduction cycle of Ginkgo biloba growing in Guizhou Province, China versus Massachusetts, USA. Location Guizhou, China (25 North latitude) Massachusetts, USA (42 North latitude) Pollination mid-March to early April mid-May Seed Abscission mid-September late October to early November Outdoor Germination mid-March mid- to late June Ginkgo Sexuality 17 mid to late spring (March through early June), while those in cold climates will not germinate until later in the summer (late June through early August), when conditions for establishment are less favorable and the seedlings have less time to accumulate carbohydrates before going into winter dormancy. In this regard, it is worth noting that in Tuo Le Village in Guizhou Province, ginkgo seeds sown outdoors would typically germinate in March, while the same seed sown outdoors in Boston did not germinate until May 29, approximately two months later. From an ecological perspective, the complex phenology of ginkgo's sexual reproduction cycle may well have constrained the species' ability to migrate, independently of humans, into cold-temperate regions with short growing seasons, and probably accounts for its A fossilized leaf of Ginkgo yimaensis. limited warm-temperate distribution and Zheng (2003), have pushed the lineage of as a wild or semi-wild tree in the mountains of G. biloba-type ovules back to the Lower Crecentral and eastern China (Li et al. 1999, Xiang taceous, about 120 million years ago. This et al. 2006, Wei et al. 2008). Table 2 presents a suggests the possibility that the seeds of G. comparison of the phenology of Ginkgo biloba's yimaensis could have possessed a temperaturesexual reproduction cycle in Guizhou Province, sensitive, developmental-delay mechanism China versus Massachusetts, USA. similar to that of G. biloba. Such a trait would Evolutionary Implications have allowed this species to reproduce successThe fossil species Ginkgo adiantoides existed in fully in regions of the northern hemisphere that the northern hemisphere from the Upper Crewere undergoing dramatic cooling after a long taceous through the Middle Miocene (roughly period of warm conditions. Indeed, Zheng and 70 to 12 million years ago) and is considered by Zhou (2004) have proposed that \"the drastic paleobotanists to be morphologically indistinclimatic changes during the Upper Jurassic and guishable from the modern G. biloba (Tralau Lower Cretaceous, around 140 to 150 million 1968). Most of the ginkgo fossils from this time years ago, were responsible for the transformaperiod in Europe and North America come from tion of the ovulate organs of the G. yimaensis sites above 40 N latitude that were originally type into the modern G. biloba type,\" includdisturbed stream margins and levee environing the development of short shoots, the reducments, and typically occurred in association tion and protection of ovulate organs, and the with a consistent set of riparian plants, includproduction of larger seeds. Ginkgo biloba's ing Cercidiphyllum, Metasequoia, Platanus, temperature-sensitive, embryo-developmentand Glyptostrobus (Royer et al. 2003). delay mechanism could well have been another Fossils of a new Ginkgo species (G. yimaenclimate-induced Cretaceous innovation--an sis) from Liaoning Province, China, recently evolutionarily primitive but ecologically funcdescribed by Chinese paleobotanists Zhou tional form of seed dormancy. NANCy ROSE 18 Arnoldia 66\/2 Ginkgo Seed Dispersal Researchers studying various ginkgo populations in Asia have reported a number of animals feeding on, and presumably dispersing, the malodorous, nutrient-rich seeds. In China, dispersal agents include two members of the order Carnivora: the leopard cat (Felis bengalensis, family Felidae) in Hubei Province and the masked palm civet (Paguma larvata, family Viveridae) in Zhejiang Province (Del Tredici et al. 1992). In Japan, where ginkgo was introduced from China some 1,200 years ago, another member of the order Carnivora, the raccoon dog (Nyctereutes procyonoides, family Canidae), has been documented feeding on ginkgo seeds, and its droppings have been found to contain intact seeds which germinated the following spring (Rothwell and Holt 1997). The existence of three reports of omnivorous members of the Carnivora consuming whole ginkgo seeds suggests that the rancid smelling sarcotesta may be attracting primarily nocturnal scavengers by mimicking the smell of rotting flesh--in essence acting as a carrion-mimic (Del Tredici et al. 1992). The fact that ginkgo seed germination percentage is enhanced by removal of the sarcotesta lends further credence to this theory. Ancient Dispersal Agents In 2002, Zhou and Zhang reported the discovery in China of a long-tailed bird (Jeholornis sp.) from the Early Cretaceous with a large number of ginkgo-like seeds in its crop. This provides direct evidence that early birds potentially could have been involved in seed dispersal activities, although the seeds' intact nature suggests they were destined for digestion in the gizzard. In general, Ginkgo biloba seeds do not fit the typical profile of a fruit dispersed by modern birds (van der Pijl 1982). Prior to the discovery of Jeholornis, most of the speculation about Cretaceous ginkgo dispersal agents centered on dinosaurs, based primarily on their temporal overlap. If dinosaurs were involved with the dispersal of ginkgo seeds, it probably would have been carrion feeding scavengers, with teeth adapted to tearing and swallowing flesh, rather than herbivores with grinding dentition that would have A spontaneous ginkgo sapling growing out of a karst rock formation at Niu Tang village in Wuchuan County, Guizhou Province. crushed the thin-shelled seeds. At any rate, any connection between dinosaurs and ginkgo seed dispersal is, at best, conjecture based on circumstantial evidence. Ginkgo's Future By rights, Ginkgo biloba should have gone extinct long ago along with all of its close relatives. The fact that it did not provides botanists with a unique window on the past--sort of like having a living dinosaur available to study. As remarkable as ginkgo's evolutionary survival is, the fact that it grows vigorously in the modern urban environment is no less dramatic. Having survived the climatic vicissitudes of the past 120 million years, ginkgo is clearly well prepared--or, more precisely, preadapted--to handle the climatic uncertainties that seem to be looming in the not too distant future. Indeed, should the human race succeed in wiping itself out over the course of the next few centuries, we can take some comfort in the knowledge that the ginkgo tree will survive. Ginkgo Sexuality 19 This ginkgo, growing as a street tree in New Brunswick, New Jersey, shows the species' outstanding yellow fall color. 20 Arnoldia 66\/2 True survivors, these severely pruned ginkgos on a Tokyo street are growing in spite of cramped planting spaces and air pollution. Ginkgo Sexuality 21 Acknowledgements The author would like to thank Elisabeth and Jim Dudley of the Highstead Arboretum in Redding, Connecticut for their generous support of a trip to Guizhou Province in 2002, Professors Shi Jikong of Guizhou University and Fu Cheng-Xin of Zhejiang University for organizing the trip, and Jianhua Li of the Arnold Arboretum for help with all aspects of the project. Literature Cited Del Tredici, P. 2000. The evolution, ecology, and cultivation of Ginkgo biloba. In Ginkgo biloba, ed. T. Vanbeek. Amsterdam: Harwood Academic Publ. Del Tredici, P. 2007. The phenology of sexual reproduction in Ginkgo biloba. The Botanical Review 73(4): 267278. Del Tredici, P., H. Ling, and y. Guang. 1992. The Ginkgos of Tian Mu Shan. Cons. Biol. 6: 202210. Fan, X. X., L. Shen, X. Zhang, X. y. Chen, and C. X. Fu. 2004. Assessing genetic diversity of Ginkgo biloba L. (Ginkgoaceae) populations from China by RAPD markers. Biochem. Gen. 42: 269278. Friedman, W. E. 1986. Photosynthesis in the female gametophyte of Ginkgo biloba. Am. J. Bot. 73: 12611266. Friedman, W. E. and E. M. Gifford. 1997. Development of the male gametophyte of Ginkgo biloba: a window into the reproductive biology of early seed plants. In Ginkgo biloba--a global treasure. eds. T. Hori, R. W. Ridge, W. Tulecke, P. Del Tredici, J. Tremouillaux-Guiller, and H. Tobe. Tokyo: Springer-Verlag. Handa, M., y. Iizuka, and N. Fujiwara. 1997. Ginkgo landscapes. In Ginkgo biloba--a global treasure. eds. T. Hori, R. W. Ridge, W. Tulecke, P. Del Tredici, J. Tremouillaux-Guiller, and H. Tobe. Tokyo: Springer-Verlag. He, S. A., y. Gu, and Z. J. Pang. 1997. Resources and prospects of Ginkgo biloba in China. In Ginkgo biloba--a global treasure. eds. T. Hori, R. W. Ridge, W. Tulecke, P. Del Tredici, J. Tremouillaux-Guiller, and H. Tobe. Tokyo: Springer-Verlag. Hirase, S. 1896. On the spermatozoid of Ginkgo biloba. Bot. Mag. (Tokyo) 10: 325328 (in Japanese). Kochibe, N. 1997. Allergic substances of Ginkgo biloba. In Ginkgo biloba--a global treasure. eds. T. Hori, R. W. Ridge, W. Tulecke, P. Del Tredici, J. Tremouillaux-Guiller, and H. Tobe. Tokyo: Springer-Verlag. Li, J.W., Z. G. Liu, y.G. Tan and M. B. Ren. 1999. Studies on the Ginkgo at Jifo Mountain. Forest Research 12: 197201 (in Chinese). Matsumoto, K., T. Ohta, M. Irasawa, and T. Nakamura. 2003. Climate change and extension of the Ginkgo biloba L. growing season in Japan. Global Change Biol. 9: 16341642. Parliment, T. 1995. Characterization of the putrid aroma compounds of Ginkgo biloba fruits. In Fruit flavors: biogenesis, characterization, and authentication. eds. R. Rouseff and M. Leahy. Am. Chem. Soc. Symp. Ser., 596. Rothwell, G. W. and B. Holt. 1997. Fossils and phenology in the evolution of Ginkgo biloba. In Ginkgo biloba--a global treasure. eds. T. Hori, R. W. Ridge, W. Tulecke, P. Del Tredici, J. Tremouillaux-Guiller, and H. Tobe. Tokyo: Springer-Verlag. Royer, D. L., L. J. Hickey, and S. L. Wing. 2003. Ecological conservatism in the \"living fossil\" Ginkgo. Paleobiology 29: 84104. Shen, L., X. y. Chen, X. Zhang, y. y. Li, C. X. Fu, and y. X. Qiu. 2005. Genetic variation of Ginkgo biloba L. (Ginkgoaceae) based on cpDNA PCR_ Rflps: inference of glacial refugia. Heredity 94: 396401. Tralau, H. 1968. Evolutionary changes in the genus Ginkgo. Lethaia 1: 63101. Van der Pijl, L. 1982. Principles of dispersal in higher plants, 3rd ed. Berlin: Springer-Verlag. Wei, G., y. X. Qui, C. Chen, Q. ye, and C. X. Fu. 2008. Glacial refugia of Ginkgo biloba L. and human impact on its genetic diversity: evidence from chloroplast DNA. J. Integrat. Plant Biol. 50(3): 368374. Xiang, B. X, Z. H. Xiang, and y. H. Xiang. 2006. Investigation of wild Ginkgo biloba in Wuchuan County of Guizhou, China. Guizhou Sci. 24: 5667 (in Chinese). Zheng, S. and Z. Zhou. 2004. A new Mesozoic Ginkgo from western Liaoning, China and its evolutionary significance. Rev. Palaeobot. and Palynol. 131: 91103. Zhou, Z. and F. C. Zhang. 2002. A long-tailed, seed-eating bird from the early Cretaceous of China. Nature 418: 405409. Zhou, Z. and S. Zheng. 2003. The missing link in Ginkgo evolution. Nature 423: 821822. Peter Del Tredici is a Senior Research Scientist at the Arnold Arboretum. A more extensive discussion of this topic can be found in the author's article \"The Phenology of Sexual Reproduction in Ginkgo biloba: Ecological and Evolutionary Implications\", 2007, The Botanical Review 73(4): 267278. "},{"has_event_date":0,"type":"arnoldia","title":"The Fruits of Autumn","article_sequence":3,"start_page":22,"end_page":27,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25447","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed14ea36b.jpg","volume":66,"issue_number":2,"year":2008,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"The Fruits of Autumn Nancy Rose utumn is prime time for observing a great array of maturing fruits on woody plants. Fleshy types like pomes, drupes, and berries are often brightly colored and highly noticeable at this time of the year. Fallfruiting trees and shrubs--viburnums (Viburnum spp.), crabapples (Malus spp.), mountain ash (Sorbus spp.), beautyberries (Callicarpa spp.), and hollies (Ilex spp.), to name a few--provide a showy display, especially as deciduous leaves begin to fall. In addition to adding color to the landscape, fall-fruiting plants also serve as an important food source for birds. Other fruiting structures seen in autumn are less showy but still interesting. Pods, samaras, and inflated capsules are some of the diverse forms to be seen. As anyone who has ever tried to learn woody plants knows, fruits often provide the key for correct identification. Here are some examples of fruits to look for this fall: ALL PHOTOS By THE AUTHOR EXCEPT AS INDICATED ROBERT MAyER A Grape honeysuckle, (Lonicera reticulata) Common persimmon (Diospyros virginiana) The word \"berry\" is often used to describe just about any rounded, juicylooking fruit, but botanically speaking a berry is a fleshy, indehiscent (not splitting open at maturity) fruit that develops from a single pistil and contains one or multiple seeds. A number of woody plants bear berries including vines like Vitis (grape), Actinidia (kiwi), and Parthenocissus (Virginia creeper, Boston ivy). Both vine and shrub species of Lonicera (honeysuckle) have berries, often attractive bright red ones. Common persimmon (Diospyros virginiana) is one of few large trees that produces true berries; look for the golden orange, globe-shaped fruits persisting on branches through late autumn. Fruits of Autumn 23 MICHAEL DOSMANN Clockwise from upper left: Donald Wyman crabapple (Malus `Donald Wyman') Korean mountain ash (Sorbus alnifolia) Chinese sand pear (Pyrus pyrifolia) Black chokeberry (Aronia melanocarpa) A pome is a fleshy, indehiscent fruit that develops from a compound ovary set within a fleshy floral cup or tube. Multiple seeds are found in the core of the fruit. Pomes are the fruits of a number of well-known genera in the rose family (Rosaceae), including Malus (apple, crabapple), Sorbus (mountain ash), Pyrus (pear), Crataegus (hawthorn), Aronia (chokeberry), Cotoneaster, and Pyracantha (firethorn). MICHAEL DOSMANN 24 Arnoldia 66\/2 Clockwise from upper left: Sapphireberry (Symplocos paniculata) American cranberrybush (Viburnum trilobum) Purple beautyberry (Callicarpa dichotoma) Winter Red winterberry (Ilex verticillata `Winter Red') Another common berrylike fruit found on woody plants is the drupe. A drupe is a fleshy, indehiscent fruit containing a single seed which is surrounded by a stony endocarp. Many of the showiest fall-fruiting shrubs and small trees bear drupes, including viburnums (Viburnum spp.), beautyberries (Callicarpa spp.), dogwoods (Cornus spp.), and hollies (Ilex spp.). Many delicious drupes are found in the genus Prunus including cherries, plums, and peaches. Fruits of Autumn 25 A hip is a pomelike structure formed by a fleshy hypanthium (a cup-shaped structure formed from fused floral parts at the flower's base) which surrounds multiple achenes (small, dry fruits containing single seeds). The term hip is used specifically for roses (Rosa spp.). The large, scarlet hips of Rosa rugosa (left) give it one of its common names: beach tomato. Aggregate fruits are composed of numerous small fruits that develop from multiple pistils in a single flower. Raspberry fruits, for example, are aggregates of drupelets. Magnolias produce conelike aggregates of follicles; at maturity, each follicle opens to reveal a seed covered by a brightly colored aril (fleshy seed coat) and attached by a stretchy thread. The fruit of a hybrid sweetbay magnolia (Magnolia virginiana) is seen here (right). MICHAEL DOSMANN Multiple fruits develop when the fruits derived from numerous individual flowers in an inflorescence fuse together to form what appears to be a single fruit. Pineapple (Ananas spp.) and mulberry (Morus spp.) are examples of multiple fruits. The unique, baseball-sized green fruits of osage orange (Maclura pomifera), shown at left, are also multiple fruits. 26 Arnoldia 66\/2 Built to be carried by the wind, samaras are winged achenes. The papery wing part of the structure takes variable forms; for example, in elms (Ulmus spp.) the wing encircles the achene, in ash (Fraxinus spp.) the wing extends like a paddle from a single achene, and maples (Acer spp.) bear paired (two-winged) samaras that usually split apart when they mature and fall. The size and wing angle of maple samaras provide a good identification key among species. Three-flowered maple (Acer triflorum) bears triplets of two-winged samaras. Another samara variation--a single achene dotted in the middle of the wing--is seen in this red-fruited form of the notoriously seedy tree-of-heaven (Ailanthus altissima f. erythrocarpa). Exclusive to oaks (Quercus spp.), acorns are hard-shelled seeds (nuts) nested in cupshaped involucres. Acorn size and degree of involucre extension on the nut provide a good clue when trying to identify oak species. Noted for their extensively fringed involucres, the acorns of bur oak (Quercus macrocarpa) are seen in this image. Fruits of Autumn 27 Many plants bear seed-holding capsules but the forms of these dry, dehiscent (splitting open at maturity) fruits vary widely. The inflated, paper-lanternlike capsules found on golden rain tree (Koelreuteria paniculata, left) turn from green to tan--sometimes with a blush of pink--and often persist well into the winter. Also shown (right) are the small, rounded capsules of summersweet (Clethra alnifolia), filled with numerous tiny seeds. Pods are dry, dehiscent or indehiscent fruits that contain seeds. The legume family (Fabaceae) is well-known for producing pods as its fruiting structure. Woody plants in this family include honey locust (Gleditsia spp.; pods of G. triacanthos pictured), Kentucky coffee tree (Gymnocladus dioicus), wisteria (Wisteria spp.), and silk-tree (Albizia julibrissin). "},{"has_event_date":0,"type":"arnoldia","title":"Book Review: Fruits and Plains: The Horticultural Transformation of America","article_sequence":4,"start_page":28,"end_page":31,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25444","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed1708928.jpg","volume":66,"issue_number":2,"year":2008,"series":null,"season":null,"authors":"Schlereth, Thomas J.","article_content":"Book Review: Fruits and Plains: The Horticultural Transformation of America Thomas J. Schlereth Fruits and Plains: The Horticultural Transformation of America Philip J. Pauly. Harvard University Press, Cambridge, Massachusetts, 2007. 336 pages. ISBN-13: 978-0-674-02663-6 M any readers, at first glance, may find this book's main title a bit puzzling. What do pomology and plains have in common? The author intends this minor mystery but he does provide several clues in his introduction and the nine chapters that follow. I must admit I had not completely grasped his full meaning until reaching his closing chapter where a complete explanation is found. Out of respect for the author's book-craft, I too will leave this resolution for the end. Long before arriving at the book's conclusion, I knew that what I was reading was a provocative and persuasive re-interpretation of several interrelated research fields; namely American plant pathology, biogeography, and cultural history. Moreover, it was a brilliant and novel re-interpretation of nineteenth-century American history using American cultivated plants as a primary resource. Beginning with the introduction (\"Taking the History of Horticulture Seriously\"), Philip J. Pauly launches his methodology of interconnecting American horticultural history with American cultural history. This fruitful hybrid yields many useful insights, one of which is how our perpetual indulgence in claiming to be exceptional in our nationhood can also be found, repeatedly, in our horticultural history. As one might expect of a cultural historian, Pauly frequently reminds us of a more universal issue evident in all of our interactions with the natural word: that is, whether we are home gardeners or plant scientists, landscape archi- tects or arboretum directors, USDA bureaucrats or environmental historians, we all culture nature. When we horti-culture nature, its plants become, to various degrees, natural artifacts subject to various forms of human artifice. Hence there are two general perspectives that characterize Pauly's achievement. First, one can see it as a revisionist interpretation in American Character historiography, a subfield in interdisciplinary American Studies scholarship since the 1950s. Second, the book is also a carefully documented survey of how Americans, despite their professed objectivity (scientific and otherwise), historically brought various types of cultural baggage (political and economic; regional and religious; profes- Book Review 29 sional and personal) to their several centuries of interactions with other living organisms and particularly with plants and plant pests. To document this dual approach--explaining both American history and the history of American horticulture--Pauly analyzes the motives and actions of a cadre of Americans who cultured nature in diverse ways and often for divergent purposes. Many will be familiar to Arnoldia readers: for instance, Thomas Jefferson, Frederick Law Olmsted (Senior and Junior), Charles Hovey, Charles S. Sargent, Horace J. McFarland, Jens Jensen, and Liberty Hyde Bailey. Also studied are less wellknown but influential plant culturists such as David Hosack, Beverly T. Galloway, William Saunders, Ephraim Bull, Charles T. Simpson, Daniel Simberloft, Charles L. Marlatt, and Katherine Bates. With these dramatis personae, Pauly explores several additional subthemes. In chapter one, for example, he stages Thomas Jefferson as an early exemplar of American horticultural chauvinism, particularly in his Notes on the State of Virginia (1785, 1787) written, in part, to answer Guillame Raynal's Historie de deux Indes (1770), a European best seller that claimed the New World's flora, fauna, climate, as well as its native peoples and even its recent emigrant Europeans were all in a state of continual anthropological and biological degeneracy. In chapter one, he also provides early definitions for terms readers will find throughout the book: first, a vocabulary of \" N-words\": nature, natural, naturalism, nationalism, and nativism; second, a litany of \"C-words\" that no cultural historian can do without: culture, cultural, and culturalism, plus related \"culture\" nomenclature that Pauly uses frequently. Chapter two initiates another important book topic--the tensions and controversies (diplomatic, military, economic, political, and scientific) that have been factors in the history of plant introductions and plant pests all arriving in increasing numbers to a supposedly virgin land. The first culprit is the Hessian fly (Mayetiola destructor) which Pauly discusses as \"America's first invasive\" as well as \"the nation's first postcolonial public scientific issue.\" This initial late eighteenth-century debate over invasives and introductions resurfaces in several places throughout the book in its survey of nineteenth- and early twentiethcentury arguments over exotic vs. native species as well as the horticultural practices (organic vs. chemical) in solving plant pathologies. Chapters five and six, cleverly named by Pauly as \"Immigrant Aid: Naturalizing Plants in the Nineteenth Century\" and \"Mixed Borders: A Political History of Plant Quarantine,\" document the local, regional, and national aspects of these prolonged conflicts, many of which are still contested issues in present-day horticulture. Massachusetts Gypsy Moth Commission employees scraping gypsy moth egg masses off of a notable elm in Malden, Massuchusetts in the early 1890s. From The Gypsy Moth (1896), Edward H. Forbush and Charles H. Fernald. 30 Arnoldia 66\/2 STEREOPTICAN IMAGE COURTESy OF WELLESLEy COLLEGE ARCHIVES In chapter six's subtitle, another Pauly interpretive emphasis appears. He recognizes that plants have politics in the sense that people culture plants with political (and other) motives. For some readers, however, his extremely detailed accounts of the political infighting among plant importers and breeders, university science faculty and nursery growers, government officials and departments as well as plant collection administrators may Book Review 31 prove too tedious a tale to stay with until the chapter's conclusion. Turning back to chapters three and four, respectively titled \"The Development of American Culture, with Special Reference to Fruit\" and \"Fixing the Accidents of American Natural History: Tree Culture and the Problem of the Prairie,\" we find major clues to the book's main title as well as nineteenth-century America's fascination with pomology. It also introduces us to Midwestern horticultural biogeography, one of the book's three such foci--the other two being the country's northeastern corridor and the anomaly of the \"horticultural construction\" of Florida. The latter history turns out also to have interesting ties to northeastern plant culturists, as diverse as diplomat Henry Perrine, proprietary town builder and citrus magnate Henry Stanford, railroad and luxury hotel entrepreneur Henry Flagler, plus the USDA's David Fairchild (after whom the Fairchild Tropical Botanic Garden in Coral Gables is named), and America's most famous nineteenth-century woman abolitionist and author, Harriet Beecher Stowe. (Interestingly, author Pauly grew up in Ohio, one gateway to--as well as an important part of--the Midwest's horticultural hearth.) Pauly's chapter nine (titled \"Culturing Nature in the Twentieth Century\") is unfortunately only a 28-page introduction to what might have been a larger Fruits and Plains or a second volume as its sequel. Here we find important developments such as the founding, at long last, of a National Arboretum in 1927, and the influential Midwestern prairie restoration by James Curtis and Aldo Leopold at the University of Wisconsin Arboretum in 1936. Also treated are the importance of the American Society for Horticultural Science and the enormous multiplication of garden clubs nationwide, plus a brief survey of \"How Pests Became Invasive Species.\" Given its brevity, the chapter is a tantalizing but selective overview of an extremely complicated and conflicted century in American horticultural history. In beginning his final chapter, Pauly references the poetry, travels, and academic career of Katherine (Kitty) Bates, an undergraduate and later a lifelong English professor at Wellesley College. Pauly muses that Bates, both as student and teacher on the Wellesley campus, could gaze across Lake Waban and see the highly cultured conifer topiary garden and arboretum at the estate of H. H. Hunnewell, one of New England's most well-known horticulturists and a generous benefactor of the Arnold Arboretum. In 1893, Professor Bates took a combined pleasure\/professional trip to teach a summer-school course at Colorado College. En route she visited Chicago's World's Columbian Exposition designed in part by Frederick Law Olmsted, Sr., travelled through Kansas prairies and wheat fields, and climbed Pikes Peak for a majestic view of the seemingly never-ending Great Plains. Atop that mountaintop, she reflected on all that she had seen on her westerning odyssey. On the peak, the beginning words of a poem also came to her. It was published in 1895 by The Congregationalist as its Fourth of July number. New yorker Samuel A. Ward set the poem to music and we have sung it ever since, a geographical and horticultural counter point to Francis Scott Key's militant navel ode whose melody Key borrowed from a British drinking song. Professor Pauly deploys Professor Bates's verses (obviously \"the fruited plain\") to announce his final chapter titled \"America, The Beautiful.\" More an epilogue than a chapter, it serves as his own anthem to his subject's meaning in both American horticultural history and American cultural history. He concludes by noting that the Bates metaphor provided him with \"a kind of professional and personal perspective\" by which to summarize and to reflect on his book's methodology (the transformation of horticulture by American culture, culturing, and culturists) and its ambitious scope and synoptic brilliance (to offer an answer, in my judgement, to the question: \"What's American about American nature?\"). In his moving, intimate acknowledgements-- placed significantly but uncharacteristically at the end of his conclusion--he alludes to his personal battle with lymphoma cancer. Phillip J. Pauly died of the disease in April, 2008, at age 57, and American historical scholarship lost one of its most insightful culturists. Thomas J. Schlereth is Professor of American Studies and History at the University of Notre Dame. "},{"has_event_date":0,"type":"arnoldia","title":"An Excerpt From Fruits and Plains: The Horticultural Transformation of America","article_sequence":5,"start_page":32,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25443","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed170856f.jpg","volume":66,"issue_number":2,"year":2008,"series":null,"season":null,"authors":"Pauly, Philip J.","article_content":"An Excerpt From Fruits and Plains: The Horticultural Transformation of America Philip J. Pauly Prairie Spirit ortheasterners' struggles to garden landscape were recapitulated, in a shorter time span and with greater seriousness, in Illinois and Wisconsin. Interest in replicating familiar Anglo-Hudson scenery competed with desires to evoke the regionally distinctive prairie. Landscape historians have focused on the pre-World War I innovations of the Danish German immigrant Chicago park designer Jens Jensen and the American horticulturist Wilhelm Miller. I suggest, however, that Jensen's and Miller's \"prairie style of landscape gardening\" drew so much from German and Olmstedian naturalism, and placed so much emphasis on shrubs and trees, that it contained little that was distinctive. The truly important development occurred, not on Chicago parklands or North Shore estates in the 1910s, but in southern Wisconsin in the 1930s, where Aldo Leopold planted a vast wildflower garden. [p. 187] N ldo Leopold, Norman Fassett, and Theodore Sperry were the developers of a real prairie style of landscape gardening. Between 1935 and 1940, they transformed about twenty-seven acres of old pasture in Dane County, Wisconsin, a few miles southwest of Madison, into a naturalistic garden of grasses and wildflowers that they called a prairie. This act of historical naming enabled them to resolve the problem faced by landscape gardeners from Downing to Miller. They planted a landscape that was distinguishable from, and an improvement upon, the common vegetation around it, but which was plausibly naturalistic. The University of Wisconsin Arboretum began as a provincial Olmstedian park project. In 1911 the private Madison Park and Pleasure Drive Association hired the young Massachusetts landscape architect John Nolen to prepare a comprehensive plan for the improvement of their city. Among Nolen's recommendations was the idea that the city and the university should emulate Boston and Harvard's partnership of the 1870s by establishing an arboretum-park on the shore of Lake Mendota, west of the city and the university campus. That suggestion went nowhere. The arboretum idea was revived in the late 1920s, however, by local boosters seeking to transform a failed suburban development on the small and marshy Lake Wingra, a few miles southwest of the city. They argued that the state and the university should fund a park, arboretum, and wildlife refuge as part of the ongoing initiative to establish a conservation professorship for Madison-based forester and game manager Aldo Leopold. The university approved this plan in 1932, appointed landscape architect William Longenecker to the position of executive Reprinted by permission of the publisher from Fruits and Plains: The Horticultural Transformation of America by Philip J. Pauly, Cambridge, Massachusetts: Harvard University Press, copyright 2007 by the President and Fellows of Harvard College. A Original Wisconsin Fruits and Plains 33 MOLLy FIFIELD MURRAy, UNIVERSITy OF WISCONSIN The Curtis Prairie at the University of Wisconsin Arboretum as it appears today. director, and asked Leopold to take on the arboretum's research directorship as one of his professorial duties. Disagreements arose immediately over issues of plant choice. Longenecker envisioned a landscape park containing systematically and ecologically ordered displays of all the perennials, shrubs, and forest trees that might prove hardy in Wisconsin. Visitors to the arboretum would be inspired to beautify their own properties, and would learn what different ornamentals and woodland trees looked like and which were worthwhile. Leopold wanted to send the visiting public a different message. He was uninterested in what he considered merely \"a `collection' of imported trees.\" Instead he wanted to show how much the state's vegetational quality had declined since the 1830s, and to provide a vision for improvement in the future. Advised by botany professor Norman Fassett, he proposed that the arboretum should be \"a reconstruction of original Wisconsin.\" It would be \"a bench mark, a datum point, in the long and laborious job of building a permanent and mutually beneficial relationship between civilized men and a civilized landscape.\" This disagreement was resolved by dividing the arboretum into areas controlled by either Longenecker or Leopold. For Leopold and Fassett, original Wisconsin was an essentially steady state, consisting of forest, wetland, and prairie, that had existed prior to Anglo-American settlement. (They passed over the major presence of Indians in Dane County during the Woodland Period, evident in the number of mounds--over one thousand, more than anywhere else in the United States.) Creating replicas of these plant communities on a few hundred acres would require a number of different kinds of effort. Sections with trees could redevelop on their own if there were fire suppression and culling of undesirable species. The right mix of wetland vegetation depended largely on 34 Arnoldia 66\/2 COURTESy UNIVERSITy OF WISCONSIN-MADISON ARCHIVES steam dredges that could change the monotonous marsh into a more varied landscape of islands and lagoons. Shoreline areas with different slopes and soil compositions could then be planted with cattails and pondweeds that would attract wildfowl. The real gardening challenge, however, was to create a \"Wisconsin prairie\" (the present-day Curtis Prairie). The basic This photograph from the 1930s shows University of Wisconsin horticultural director William Longenecker directing Civilian Conservation Corps workers prerequisite was labor. planting prairie sod. In 1934 the arboretum received a windfall when the state established a work relief camp for transients on its grounds. Then, when complaints arose about the behavior of these migrants and hoboes, the university persuaded the National Park Service to take over the camp and use it for the Civilian Conservation Corps (CCC) (see Figure 7.9). The CCC recruited a more tractable pool of young local men, and its involvement enabled the university to hire the young National Park Service plant ecologist Theodore Sperry as foreman. \"Camp Madison\" averaged about two hundred residents during the second half of the 1930s, at a cost to the federal government of more than two million dollars. The first step in the creation of a Wisconsin prairie park was to clear existing old-field growth. Tree control was a straightforward matter of destroying saplings, but was complicated by Fassett and Sperry's interest in leaving one large tree standing to evoke early settler accounts of \"oak openings\"; each year laborers had to pull up a crop of squirreland bird-distributed oak seedlings. The major problem was quack grass. Sperry and his workers sought to eliminate this Old World pasture mainstay and agricultural weed by plowing deeply, harrowing to dry out the rhizomes, and then replanting with clover to smother remaining growth. Irritating plants such as nettles and thistles were also a concern, without regard to their geographic origin. Finally, Leopold sought to suppress high-density populations (\"thickets\") of plants that were too common, such as goldenrods and asters. Once the ground was cleared, the major issues involved plant choice. In principle, Fassett and A Civilian Conservation Corps worker displays a Sperry's palette could include any of the species massive Silphium taproot. UNIVERSITy OF WISCONSIN-MADISON ARBORETUM ARCHIVES Fruits and Plains 35 NANCy ROSE associated with prairies in or near Wisconsin during the previous century. A present-day list of such plants totals between 340 and 550. But prairie gardeners in the 1930s were neither capable of nor interested in cultivating such a diverse flora. Sperry's planting list from 1935 to 1939 consisted of about fifty species. In both his exclusions and featured species, his goal was to plant an assemblage that would not be confused with common or despised pasture. The largest category of excluded species consisted of the dozens of plants that were small, had inconspicuous flowers, or were visually generic. There was minimal interest in devoting labor and space to vegetation that added little to the field's visual composition. More straightforwardly, Sperry did not replant the nettles and thistles that had been removed when the land was cleared, nor did he introduce additional species with similar properties. While some of the more memorable native species that people encountered on Wisconsin prairies were greenbrier (Smilax lasioneura), prickly Compass plant (Silphium laciniatum) was one of the forbs selected by plant ecologist Theodore Sperry for pear (Opuntia macrorhiza), and poison ivy (Toxico- the Wisconsin prairie park. dendron radicans), they were not part of the arboretum plantings. The most interesting group of exclusions was of species poisonous to livestock. Prairie larkspur (Delphinium carolinianum subsp. virescens), sundial lupine (Lupinus perennis), and death camas (Zigadenus elegans) were all visually impressive Wisconsin natives. But the prosperous rural citizens whose sensibilities Leopold wanted to touch would not have appreciated a field filled with seed-bearing specimens of the weeds they had worked for a century to eradicate. Sperry wisely emphasized familiar species that would, under proper cultivation, provide a spectacular mass display. His most frequently planted species was turkeyfoot grass (Andropogon gerardii, now commonly called big bluestem). The mostplanted forbs were stiff sunflower (Helianthus rigidus) and three species of Silphium (including compass plant and rosinweed). Others included blazing star (Liatris), prairie goldenrod (Solidago rigida), prairie rose (Rosa carolina), prairie bush clover (Lespedeza capitata), prairie coneflower (Lepachys pinnata), and prairie painted cup (Castilleja sessiliflora). They were either large (big bluestem, compass plant, and stiff sunflower could all grow ten feet high in a good summer), had conspicuous flowers (blazing star, rose, coneflower), or unusual characteristics (indicated in names such as compass plant and painted cup). While Wisconsinites might know these plants, they would have seen them only in small populations or in fields browsed by livestock. At the arboretum, by contrast, they were able to display their capabilities and to reinforce each other visually as elements of a multiacre garden. People who visited this landscape, especially in the peak summer vacation months of July and August, would experience a wonderful wildflower garden in the style of a prairie. It was both easy and pleasant to imagine that this was original Wisconsin. [pp. 190 to 194] "},{"has_event_date":0,"type":"arnoldia","title":"Collecting Sweetgum in the Wilds of Missouri","article_sequence":6,"start_page":36,"end_page":36,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25445","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed170896d.jpg","volume":66,"issue_number":2,"year":2008,"series":null,"season":null,"authors":"Alexander III, John H.","article_content":"Collecting Sweetgum in the Wilds of Missouri John H. Alexander III O n a sunny day in December, 1979, in the countryside near Zalma, Missouri, a tractor dutifully worked the upper end of a 40-acre field. In search of native tree seeds, Arboretum horticulturist Gary Koller and I were about half way across the lower end of the field when the tractor turned toward us. The farmer, clearly silhouetted against the skyline, picked up a rifle. It suddenly became clear that we were trespassing. Gary turned to me and said \"Do you have a business card?\" I don't recall the expletives or the suggestions I proffered, but I handed him a card. He ran toward the tractor, waving the card like a tiny white flag. Dumbfounded, I stood and watched, then followed. As it turned out, the farmer's brother had been shot (not fatally) by errant hunters while working in these fields a week or two earlier. The farmer was friendly, apologized for the rifle, and welcomed us to collect seeds in an uncultivated area by the river. It was there in the floodplain of the Castor River that Gary and I collected the seed from which the sweetgum (Liquidambar styraciflua, accession 1248-79-B) pictured at right was grown. At 28 years old, this sweetgum is 36 feet (11 meters) tall with a DBH (diameter at breast height) of 16 inches (40 centimeters). Typical mature height for sweetgum is around 60 to 80 feet (18 to 24 meters). Sweetgum has a pyramidal habit when young; older trees often have a rounded canopy. Its star-shaped leaves can develop striking fall color in shades of yellow, red, orange, and purple. The spiky, 1 to 1 inch (25 to 38 millimeters) diameter fruits may be dried and used in decorations, but in large numbers can be an inconvenience when they fall on lawns and walkways. Sweetgum's branch texture is variable from tree to tree; branches may be fairly smooth or have corky wings. The latter trait is impressively displayed on specimen 1248-79-B; its eye-catching abundance of large, corky, winged protrusions gives the tree great textural interest, especially in the winter. Native Ground The Arnold Arboretum is well-known for its international plant explorations, especially in China. Woody plants from around the world fill the Arboretum's collections. But collecting from wild populations of native North American plants is also important to the Arboretum's mission. Gary and I were in Missouri to attend a plant propagators' conference in St. Louis, but we had also scheduled a couple of extra days for collecting in the area. Our goal in southeast Missouri was to find species that were native to southern regions of the United States but were growing wild in a climate that was similar to our own in Boston. Sweetgum's principal native range extends from New Jersey to southern Illinois, south to eastern Texas and northern Florida. It is usually listed as hardy to USDA Zone 5, but specimens grown from seed sources in the southern part of its range may suffer significant damage in northern winters. We must have succeeded in collecting from an appropriate location--all three specimens of accession 1248-79 are in good condition. Currently, these are the only sweetgum trees in the Arboretum that are from a known wild source. In addition to sweetgum, we collected a number of other species in Missouri and neighboring Illinois, including Ohio buckeye (Aesculus glabra), pawpaw (Asimina triloba), sycamore (Platanus occidentalis), possumhaw (Ilex decidua), buttonbush (Cephalanthus occidentalis), American hornbeam (Carpinus caroliniana), and river birch (Betula nigra). These species are all fairly common, but what's important is that our collections provide a genetic representation of each of these species as it exists in the wild. When one of these plants, like sweetgum specimen 1248-79-B, turns out to have ornamental characteristics that appeal to us as gardeners, that's icing on the cake. John H. Alexander III is Plant Propagator at the Arnold Arboretum. "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23410","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd25e8928.jpg","title":"2008-66-2","volume":66,"issue_number":2,"year":2008,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"The Chinese Parrotia: A Sibling Species of the Persian Parrotia","article_sequence":1,"start_page":2,"end_page":9,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25442","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed170816b.jpg","volume":66,"issue_number":1,"year":2008,"series":null,"season":null,"authors":"Jianhua, Li; Del Tredici, Peter","article_content":"The Chinese Parrotia: A Sibling Species of the Persian Parrotia Jianhua Li and Peter Del Tredici he Persian ironwood (Parrotia persica) has a well-deserved reputation as a beautiful garden plant--mainly because of its exfoliating bark and gorgeous fall color--but also as a tough species that tolerates drought, heat, wind, and cold (Dirr 1998). Less well known is the fact that Persian ironwood has a sister species, the Chinese ironwood (Parrotia subaequalis) (Figure 1), growing about 5600 kilometers (3500 miles) away in eastern China. Remarkably, this species was correctly identified only sixteen years ago (Deng et al. 1992a). T The Persian and Chinese ironwoods are members of the witch hazel family (Hamamelidaceae), and in order to appreciate their uniqueness and evolutionary history we need to first examine one of their more familiar relatives, the witch hazels (Hamamelis). There are five species of witch hazel distributed throughout the temperate regions: H. mollis in eastern China, H. japonica in Japan, and H. virginiana, H. vernalis, H. mexicana in North America. The genus shows the intercontinental disjunct distribution between eastern Asia and North Figure 1. Geographic distribution of Parrotia persica (in green) and P. subaequalis (in red). Note that the scale bar is 400 kilometers. Parrotia 3 Hamamelis virginiana ..... 55 84 Distylium racemosum Parrotia persica ........... 91 7.83.8 mya Parrotia subaequalis ................................. 50 mya Parrotiopsis jacquemontana ......... Fothergilla major .... 10 changes Figure 2. Evolutionary relationships of Hamamelis and petalless genera, showing shift (the arrow) from insect to wind pollination. Black dot indicates the fossil calibration point and the red dot shows the divergence time of the two Parrotia species. The scale bar represents ten changes in nucleotide composition as measured along the horizontal branches of this phylogenetic tree. Changes in nucleotide composition indicate genetic evolution over time. The numbers that appear over several of the branches indicate the percentages of statistical support for those groupings. Higher numbers indicate stronger evidence of support. (mya=million years ago) America that has fascinated many scientists since the time of Asa Gray (Gray 1846). Witch hazels have four ribbonlike petals (Figure 2) that come in a variety of colors from yellow to reddish copper. Six other genera in the witch hazel family have similar ribbonlike petals and occur in Southeast Asia, Africa, Madagascar, and northeastern Australia. These genera have traditionally been considered closely related to one another and to Hamamelis because they have the same number of similarly shaped petals. But recent DNA analysis has determined that the genera with four ribbonlike petals do not form a closely related natural group because they are positioned on different branches in the MOBOT 100 Sycopsis sinensis ................... UBC Botanical Garden Distyliopsis tutcheri 4 Arnoldia 66\/1 the only known species in the genus. In 1960 Professor H. T. Chang of Sun Yat-sen University described a new species of Hamamelis--H. subaequalis-- based on a fruiting specimen that had been collected twentyfive years earlier from Yixing county of Jiangsu province, China. Its main distinguishing feature was that it produced much smaller leaves than the Chinese witch hazel (H. mollis) (Figure 3) (Chang 1960). The fact that the plant described as H. subaequalis was not recollected until 1988--some 53 years after its initial collection--led to speculation that Figure 3. The foliage of a specimen of Parrotia subaequalis growing at the Nanjing the plant had gone extinct in Botanical Garden. the intervening years. witch hazel family tree. Interestingly, in each In the fall of 1988, Miaobin Deng and colbranch of this family tree the most advanced leagues at Jiangsu Institute of Botany disgenera are those that have lost their petals, a covered a natural fruiting population of H. trait that is generally believed to correlate with subaequalis in the town of Yixing. After three the transition from insect to wind pollination years of continually monitoring the population, (Li et al. 1999). During this evolutionary transitheir patience was rewarded when the plants tion period, a few genera in the Hamamelis-- finally flowered again (Deng et al. 1992b). At Parrotia lineage developed showy parts other that point it became clear that H. subaequalis than petals with which to attract insect pollacked petals, making it dramatically different linators. For example, Parrotiopsis of the westfrom H. mollis (Figure 4). They proposed a new ern Himalayas possesses showy leaflike bracts genus--Shaniodendron--to accommodate the beneath the inflorescences, while Fothergilla species which they named S. subaequale (Deng species in the eastern U.S. have conspicuous et al. 1992a). Dr. Riming Hao, who studied the white stamen filaments (Figure 2). In contrast, floral morphology of Shaniodendron, pointed Parrotia flowers lack not only petals but also out that Shaniodendron subaequale was quite showy bracts and stamen filaments. Instead, similar to Parrotia persica, but he did not place their anthers are elongated, a characteristic it within the genus Parrotia (Hao et al. 1996). In common to wind-pollinated species including 1996, Dr. Yinlong Qiu sent some DNA of Shathe most advanced genera in Hamamelidaceae. niodendron to Jianhua Li, then a PhD candidate Thus, the shift from insect to wind pollination at the University of New Hampshire working is complete in the evolutionary branch leading on the systematics of the witch hazel family. to Parrotia, Sycopsis, Distyliopsis, and DistyHe obtained nuclear DNA sequence data from lium (Figure 2). the sample and, after comparing it with other genera of the family, determined that ShaTaxonomic History of the Chinese Parrotia niodendron was a sibling species to Parrotia The first recorded species of Parrotia--P. persica (Li et al. 1997). After seeing the DNA persica--was described by C. A. Meyer in 1831 results, Hao used this evidence to propose the and named in honor of F. W. Parrot, a German merger of Shaniodendron with Parrotia (Hao naturalist and traveler. For a long time it was and Wei 1998). Nevertheless, it seems that this P. DEL TREDICI Parrotia 5 JIANHUA LI Figure 4. The flowers of Parrotia subaequalis as shown on a sign posted at the Yixing Caves Scenic Area. treatment may take some time for people to accept since recent studies continue to use the name Shaniodendron subaequale (Fang et al. 2004; Huang et al. 2005), despite the fact that the plant is listed as Parrotia subaequalis in the Flora of China. Parrotia persica and P. subaequalis are very similar from growth habit to morphology. Both trees display exfoliating bark, have obovate leaves with bluntly toothed margins, and grow in moist habitats along streams. They bear four to seven flowers clustered in a head inflorescence subtended by broadly ovate, brownish bracts. Each flower has five sepals but no petals and four to fifteen stamens with long anthers (Figure 4). Their fruits are woody capsules consisting of two chambers, each with two brown seeds (Figure 5). Parrotia subaequalis can be easily distinguished from P. persica by its lanceolate stipules and sepals fused into a shallow saucer-shaped calyx (Hao et al. 1996). When did Parrotia persica and P. subaequalis diverge? Recent DNA work in Jianhua Li's laboratory has shown that witch hazels (Hamamelis) are more primitive than the petalless genera in Hamamelidaceae. The evolutionary sequence of the petalless genera appears in the order of Fothergilla, Parrotiopsis, Parrotia, Sycopsis, and Distyliopsis plus Distylium, and the two species of Parrotia are grouped together (Figure 2). Fossils can provide evidence for the minimum age of the lineage to which they belong. Unfortunately, fossil information is often unavailable for a specific taxon. Nevertheless, if DNA molecules evolve at a constant rate, that is, a certain number of nucleotide changes per million years, we can use the total number of changes between the two species to estimate how long ago they diverged. Our statistical tests indicated that the evolution of the nuclear genes we have used to reconstruct the evolutionary history of these genera followed a clockwise manner. The next thing we needed was to calibrate the ticking rate of the molecular clock using one or more known fossil dates. Luckily, Radtke et al. (2005) found a fossil leaf that could be unequivocally assigned to Fothergilla, specifically F. malloryi. This fossil leaf is part of the Republic Flora of northeastern Washington State, dating to the late Eocene (about 50 million years ago), and thus provides a minimum separation age of Fothergilla from the branch leading to other genera (Figure 2). Based on the molecular clock calibrated using the fossil, our estimates suggest that the two species of Parrotia diverged around 7.5 million (plus or minus 3.8 million) years ago, during the Lower Miocene. This divergence time is consistent with the geological evidence JIANHUA LI Figure 5. Fruit and seed of Parrotia subaequalis. 6 Arnoldia 66\/1 P. DEL TREDICI Figure 6. Parrotia subaequalis cultivated as penjing at the Nanjing Botanical Garden. production, serious habitat competition from bamboos, and increasing human activities. It is essential to take immediate action and institute stricter measures to protect the species. Peter Del Tredici first saw two plants of Parrotia subaequalis on October 8, 1994. They were being cultivated in containers as penjing (bonsai) in a lath-house at the Nanjing Botanical Garden. At that time, the foliage had turned a beautiful, rich, deep red (Figure 6). According to the Director of the Garden, Professor Shan-an He, the plants had been collected in Jiangsu province at the Yixing Caves Scenic Area, which is located about 120 kilometers (75 P. DEL TREDICI that the cooling temperature in the Lower Miocene plus the uplifting of the Himalayas and the mountains of western China from 55 million years ago to the Middle Miocene may have restricted biological exchanges between central Asia and eastern China (Yin and Harrison 2000; Sun and Wang 2005). Forests in the Caspian region of central Asia and those in eastern Asia are both relicts of the widespread Tertiary vegetation (Wolfe 1975; Hosseini 2003; Sun and Wang 2005). Besides Parrotia, the two regions share many other woody plant genera including Acer, Albizia, Buxus, Castanea, Carpinus, Diospyros, Fagus, Pterocarya, Quercus, Sorbus, Taxus, and Zelkova. From an evolutionary and biogeographical standpoint it would be interesting to determine whether central Asian species within these genera are siblings of the eastern Asian species, and if so, whether their separation time agrees with that between the two Parrotia species. Parrotia subaequalis in China According to Chengxin Fu, Riming Hao, and various accounts in the literature, there are five populations of Parrotia subaequalis in eastern China: two each in Jiangsu and Zhejiang provinces (Huang et al. 2005) and one in Anhui (Shao and Fang 2004). Professor Fu's team is currently conducting a survey to determine the levels and patterns of the genetic diversity in Chinese Parrotia populations. The results will provide a scientific foundation for designing conservation strategies. Regeneration of Parrotia subaequalis populations will be challenging because of the species' alternate-year fruit Figure 7. Dr. Hao Riming of the Nanjing Botanical Garden with a plant of Parrotia subaequalis grown from a cutting. Parrotia 7 JIANHUA LI miles) southwest of Nanjing on the east side of Tai Lake. Both specimens had massive trunks and the larger of the two was about 50 centimeters (20 inches) tall by 70 centimeters (28 inches) across. The form of their trunks, along with their extensive yet well-healed wounds, suggested that both plants were very old. When Peter returned to the Nanjing Botanical Garden in September of 1997, he didn't see the penjing specimens but saw one young plant--recently propagated from a cutting and about 2 meters (6.6 feet) tall--growing out on the grounds of the garden (Figure 7). On September 1, 2004, we [Del Tredici and Li] had the good fortune to be able to visit the Yixing Caves Scenic Area (known as Shan Juan Park) with Professor Cheng-xin Fu and Yingxiong Qiu of Zhejiang University. Upon entering the park, the group immediately encountered a large specimen of Parrotia subaequalis growing on a steep slope above a small pond at the mouth of the largest of the karst caves. The plant was hard to miss because it was identified with a large sign with a close-up color photograph of the plant in bloom (Figure 4). The tree, which was about 6 meters (20 feet) tall, had two main trunks, the largest of which was 24 centimeters (9.4 inches) in diameter (Figure 8). The bark appeared to be at the peak of its exfoliation, with patches of fresh greenish white bark showing where sections of the old bark had sloughed off. There were no fruits on the plant--the species typically flowers only every other year--but there were numerous seedlings growing beneath it. A second large specimen was spotted about 30 meters (100 feet) away, on a slope in a mixed woodland with bamboo and other trees. We observed at least two cases where the exposed roots of this plant were producing vigorous young suckers, a phenomenon which had not been reported in the literature (Figure 9). Interestingly, sprouting from the base of the trunk was not observed on any of the trees. Later that afternoon, the group drove to Longwang Shan in Anji Xian, in northern Zhejiang Province, about 90 kilometers (56 miles) south of the Yixing Caves. This relatively small mountain is considered part of the larger Tian Mu Shan range that forms the border with Figure 8. The trunk and foliage of a Parrotia subaequalis specimen growing at the Yixing Caves Scenic Area. P. DEL TREDICI Figure 9. Root suckers from a mature specimen of Parrotia subaequalis at the Yixing Caves Scenic Area. 8 Arnoldia 66\/1 P. DEL TREDICI bark was not exfoliating as dramatically as the larger plant. Neither was producing any sprouts from the base of its trunk or any root suckers. Unfortunately there were no fruits on either plant, although there were curious hard, round, gall-like structures about a centimeter or so in diameter on many of the leaves of the smaller, double-trunked plant. Some of the notable associates growing with Parrotia subaequalis on Longwang Shan were Fortunearia fortunei, Styrax confusus, Pterostyrax corymbosum, Cornus controversa, Stewartia rostrata, and Stewartia sinensis. We were told that the Parrotia subaequalis population at Longwang Shan consisted of about twenty individuals at that time. P. DEL TREDICI Figure 10. Parrotia subaequalis on Longwan Shan, 9.5 meters (31 feet) tall with a diameter at breast height of 38 centimeters (15 inches). Anhui Province. After spending the night in comfortable accommodations at the research station, we hiked partway up the mountain to about 650 meters (2,130 feet) elevation and located two specimens of Parrotia subaequalis growing near the side of a stream, amidst a pile of boulders. The larger of the two trees was about 9.5 meters (31 feet) tall with a trunk diameter at breast height of 38 centimeters (15 inches) (Figure 10). Its bark was exfoliating in a dramatic way--shedding jigsaw-puzzle-shaped plates of old, blackish brown bark to expose conspicuous patches of greenish white bark below (Figure 11). The second specimen had a double trunk, was about 8 meters (26 feet) tall, and its Figure 11. This specimen of Parrotia subaequalis (same plant seen in Figure 10) shows a very knobby trunk, indicating that it has lost many lower branches over time. Parrotia 9 Parrotia subaequalis at the Arnold Arboretum The Arnold Arboretum has two established plants of Parrotia subequalis. So far, both of them have survived two winters outdoors and they are now about 1.5 meters (5 feet) tall. On June 23, 2005, during their first growing season at the Arboretum, seven cuttings between 5 and 10 centimeters (2 to 4 inches) long were taken from the two plants. A month later, on July 25, another nine cuttings were taken from the plants. All sixteen cuttings were treated with a five-second dip in an aqueous solution of 5,000 parts per million KIBA, stuck in flats filled with a mix consisting of half sand and half perlite, and placed in the high-humidity greenhouse under intermittent mist and fog. Remarkably, all sixteen of the cuttings rooted and three of them are planted in the nursery. With five plants now growing outdoors, the Arboretum is in a position to begin evaluating the horticultural potential of Parrotia subequalis. Successful establishment at the Arboretum also facilitates continued research on the genetics, physiology, reproductive biology, and conservation of this rare and evolutionarily important species. Acknowledgments We thank Cheng-xin Fu and Ying-xiong Qiu of Zhejiang University for their field assistance. Literature Cited Chang, H. T. 1960. Hamamelis subaequalis. Journal of Sun Yat-sen University 1: 35. Deng, M. B., H. T. Wei, and X. Q. Wang. 1992a. Shaniodendron, a new genus of Hamamelidoideae from China. Acta Phytotaxonomica Sinica 30: 5761. Deng, M. B., H. T. Wei, X. Q. Wang, P. Su, and J. Y. Xing. 1992b. On the significance of the discovery of Fothergilleae in China. Journal of Plant Resources and Environment 1: 3035. Dirr, M. A. 1998. Manual of Woody Landscape Plants. Stipes Publishing, Champaign, IL Fang, S., J. Yan, Q. Weng, and S. Huang. 2004. Population ecological status quo and protection study on Shaniodendron subaequale M. B. Deng, H. T. Wei in Longchi mountain reserve in Yixing. Journal of Jiangsu Forestry Science and Technology 31: 46. Gray, A. 1846. Analogy between the flora of Japan and that of the United States. American Journal of Science and Arts, ser. 22: 135136. Hao, R. M., H. T. Wei, and W. G. Liu. 1996. Floral morpholgy of Shaniodendron (Hamamelidaceae) and its taxonomic significance. Journal of Plant Resources and Environment 1(1): 3035. Hao, R. M. and H. T. Wei. 1998. A new combination in Hamamelidaceae. Acta Phytotaxonomica Sinica 36: 80. Hosseini, S. M. 2003. Incomparable roles of Caspian forests: Heritage of humankind. Forest Science 3: 3140. Huang, S., Y. Fang, Y. Peng, J. Yan, and S. Fang. 2005. The niche study of Shaniodendron subaequale population of Longchi mountain. Journal of Central South Forestry University 25: 8083. Li, J., A. L. Bogle, A. S. Klein, and K. Pan. 1997. Close relationship between Shaniodendron and Parrotia (Hamamelidaceae), evidence from its sequences of nuclear ribosomal DNA. Acta Phytotaxonomica Sinica 35: 481493. Li, J., A. L. Bogle, and A. S. Klein. 1999. Phylogenetic relationships of the Hamamelidaceae inferred from sequences of internal transcribed spacers (ITS) of nuclear ribosomal DNA. American Journal of Botany 86: 10271037. Radtke, M. G., K. B. Pigg, and W. C. Wehr. 2005. Fossil Corylopsis and Fothergilla leaves (Hamamelidaceae) from the lower Eocene flora of republic, Washington, U.S.A., and their evolutionary and biogeographic significance. International Journal of Plant Science 166: 347356. Shao, X. F. and G. F. Fang. 2004. Habitat survey and ex situ conservation of Shaniodendron subaequale. Journal of Anhui Forest Science and Technology 2: 1213. Sun, X. and P. X. Wang. 2005. How old is the Asian monsoon system?--Palaeobotanical records from China. Paleogeography Paleoclimatology Palaeoecology 222: 181222. Wolfe, J. A. 1975. Some aspects of plant geography of the Northern Hemisphere during the late Cretaceous and Tertiary. Annals of the Missouri Botanical Garden 62: 264279. Yin, A. and T. M. Harrison. 2000. Geologic evolution of the Himalayan-Tibetan orogen. Annual Review of Earth and Planetary Sciences 28: 211280. Jianhua Li and Peter Del Tredici are senior research scientists at the Arnold Arboretum. "},{"has_event_date":0,"type":"arnoldia","title":"Curatorial Notes: An Updated Living Collections Policy at the Arnold Arboretum","article_sequence":2,"start_page":10,"end_page":21,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25438","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed170b76f.jpg","volume":66,"issue_number":1,"year":2008,"series":null,"season":null,"authors":"Dosmann, Michael S.","article_content":"Curatorial Notes: An Updated Living Collections Policy at the Arnold Arboretum Michael S. Dosmann M useums, by definition, collect things, and in the case of botanical gardens and arboreta, those things are plants. In this quest to collect, curators must exercise discipline and prudence in determining what new things to acquire as well as which ones to remove. Garden collections can be notably challenging to curate because of the overwhelming breadth of possible biodiversity to accumulate. Thus, it is essential for curators to make use of a collections policy--a tool which defines the scope of the collection. The collections policy is mission-driven; it defines short- and long-term goals and establishes the direction of collection building. While the specifics of what items to collect may occasionally be included in the collections policy, they are typically outlined separately in a detailed collections development plan. Likewise, the tactics of curation, such as the means of acquisition, intricacies of database management, standardization of nomenclature, or tasks related to plant maintenance, are best housed within a separate procedural manual. Collections policy history at the Arboretum Since its inception, the Arboretum has built its living collection of plants with the aid of a collections policy, although the policy's content and application have varied considerably over the years. The indenture signed by the President and Fellows of Harvard College and the Trustees of Mr. James Arnold on the 29th of March, 1872, included the original collections policy: \"The Arnold Arboretum... shall contain, as far as is practicable, all the trees, shrubs, and herbaceous plants, either indigenous or exotic, which can be raised in the open air at the said West Roxbury...\" It was simple, direct--and too broad for Charles S. Sargent, the Arboretum's first curator and director. While he followed the spirit of the indenture's charge with aplomb, acquiring as many taxa as possible, the focus quickly shifted almost solely to woody plants, leaving most herbaceous plants out of the permanent collections. His keen interest in the floras of North America and eastern Asia, no doubt influenced by his mentor Asa Gray, led to substantive biogeographic collections from these locales. And Sargent's fascination with ornamentals resulted in the acquisition of many horticultural plants, including great numbers of botanical formae and varieties that are now considered cultivars. Sargent (1922) estimated that during the Arboretum's first half-century some 6,000 taxa grew in the collections. But space became limited in the 265-acre landscape, and the collections became crowded. The problem became acute in the years following Sargent's death in 1927; in the absence of his careful direction the collections multiplied unchecked. To respond to this dilemma, landscape architect Beatrix Farrand was hired in 1946 by Karl Sax, Arboretum director at the time, to create a restorative plan. In her assessment (Farrand 1946), she questioned whether \"the comparatively small acreage of the Arboretum can wisely accommodate all the species and varieties of woody plants of the temperate regions.\" The recommendation that she and the Administration came up with was that the collections grown in Jamaica Plain would be \"the best and most ornamental\"; research plants that lacked the desired showiness, yet had scientific merit, would be transferred to the Case Estates in Weston, where they could be lined-out in experimental nurseries. This strategy's execution was left to Donald Wyman, the Arboretum's horticulturist. Wyman undoubtedly sighed in relief with this decision. He acknowledged (Wyman 1947) the difficulty of maintaining an expanding number of plants solely at Jamaica Plain given the institutional reticence to modify any collections following Sargent's death (particularly those that Sargent had a hand in building). In theory, this splitsite solution allowed the living collections to Living Collections Policy 11 HERBERT W. GLEASON, ARCHIVES OF THE ARNOLD ARBORETUM Arboretum planting space was already filling up when this photograph was made in May of 1930, a few years after Sargent's death. The photo shows Korean azalea (Rhododendron yedoense var. poukhanense) and other plants on Bussey Hill. 12 Arnoldia 66\/1 PHOTOS BY MICHAEL S. DOSMANN Starting in the late 1970s, the Arboretum shifted its priority to collecting plant material of documented wild origin. Representative plants collected on expeditions made during this period include (clockwise from upper left): Weigela subsessilis collected in the Republic of Korea in 1977, Sorbus yuana collected in the People's Republic of China during the Sino-American Botanical Expedition in 1980, and Cotinus coggygria from the 1980 expedition to the Russian Federation (then the U.S.S.R.). Living Collections Policy 13 remain comprehensive--as per the original collections policy of 1872--while providing focus to the two sites: research collections in Weston and ornamental collections in Jamaica Plain. For the next 30 years, this practice continued and the Arboretum landscape in Jamaica Plain accrued great numbers of ornamental taxa, particularly cultivars under evaluation. This swing was reinforced by the post-war proliferation of cultivars introduced by the nursery industry, the institutional goal of becoming a showcase of horticultural material, and practically complete cessation of plant exploration efforts. It is important to note that while the collections policy did not shift, per se, its method of realization did. In the late 1970s, a shift again took place-- this time with an eye towards documentation, the prime metric used to assess a collection's value. While material of cultivated origin may carry with it notable documentation, its value is generally eclipsed by material of wild origin, particularly once it has been verified to identity. Thus, in a new living collections policy, priority shifted away from ornamental and toward botanical taxa (Spongberg 1979). While the emphasis was placed upon botanical taxa of wild origin, provisions were in place to accession or maintain garden-origin plants (as temporary placeholders), as well as cultivars--provided they were of historic significance (i.e., those with Latinized epithets proposed prior to 1953). This policy change coincided with the reinstated tradition of field collection of germplasm, both domestically and abroad. As a result, many new acquisitions of documented wild origin again crossed the Arboretum's threshold, particularly in Jamaica Plain. With respect to the practice of growing material in both Jamaica Plain and Weston, Peter Ashton (1979) reflected that the two-site strategy had come at a cost: the loss of valuable germplasm which did not survive the transfer from Jamaica Plain to Weston, including original introductions of species by E. H. Wilson and other explorers. The ambitious goal of acquiring everything--maintained in two separate sites--was too lofty, particularly with the resources available, and Jamaica Plain was deemed the primary repository. This formal policy direction was sustained for the next decade, and then reaffirmed in 1991 (Liv- ing Collections Long-Range Planning Committee 1991). As in the 1979 version, the goal stated that \"the living collections of the Arnold Arboretum were to consist of a scientific collection of entities tied to botanical, not horticultural nomenclature.\" Because the emphasis was placed on names and not necessarily taxonomy, a great deal of space in the new policy was dedicated to the \"problem of cultivars and their relationship to taxa of infraspecific botanical rank.\" The need for a collections policy update Shortly after joining the staff as Curator of Living Collections in January of 2007, I convened the Living Collections Committee to review the Arboretum's existing living collections policy and place it in context with current, as well as future, institutional needs. After thorough discussion and assessment, we restructured the policy with several broad goals in mind: The policy needs to describe the entire scope of our living collection, including collections that previously had not been highlighted such as the Larz Anderson Bonsai Collection and plants in our natural areas. It should also articulate levels of commitment, or priority (i.e., high to low), depending upon the type of collection. This would allow us greater flexibility as well as focus in collections development. The policy should not perpetuate the hierarchy between wild-origin and cultivated material. Instead, the emphasis should be placed on the level of documentation associated with individual accessions, as well as their programmatic use(s) in furthering the mission of the institution. This is particularly important when we consider the immense research potential of the collections (Dosmann 2007). The policy must be clear and usable, yet not burdened by too many details; the policy was not intended to be a procedural manual. Instead, we appended it with a list of operational definitions to aid in interpretation. Here is the result: the current living collections policy for the Arnold Arboretum. Notice that in spirit, it has remained true to the original plan of 1872; additional details have been added for clarity and for establishing organization and a sense of priority. Interspersed within the official policy below are text boxes and figures that provide illustrative examples and additional information. 14 Arnoldia 66\/1 Living Collections Policy Policy reviewed and approved on 10 September, 2007 MISSION STATEMENT The Arnold Arboretum of Harvard University discovers and disseminates knowledge of the plant kingdom to foster greater understanding, appreciation, and stewardship of the Earth's botanical diversity and its essential value to humankind. I. INTRODUCTION A. PURPOSE OF THE LIVING COLLECTIONS POLICY The Living Collections Policy of the Arnold Arboretum guides the development, management, and enhancement of the institution's Living Collections, and applies to all plants outlined below under Scope of the Living Collections. The Living Collections Policy is written and administered by the Living Collections Committee, which comprises the Curator of Living Collections (Chair of the Committee), Deputy Director, Manager of Horticulture, Manager of Plant Records, Manager of the Dana Greenhouses and Nursery, and Senior Research Scientist; it is further reviewed and approved by the Director. The Living Collections Policy is reviewed every five years and revised as needed. Operational procedures related to implementation of this and related policies are detailed in the Arboretum's General Procedures for Managing the Flow of Plants through the Department of Horticulture (January 2007). B. PURPOSE OF THE LIVING COLLECTIONS The Living Collections of the Arnold Arboretum are essential to achieving its mission as a research institution dedicated to improving the understanding, appreciation, and preservation of woody plants. As a national and international resource for research in the various fields of plant biology and beyond, the Arboretum's Living Collections are actively developed and managed to support scientific investigation and study, as well as key educational and amenity roles. C. LEGAL AND ETHICAL CONSIDERATIONS Activities related to the development, management, and use of the Arnold Arboretum's living collection comply with all relevant local, state, federal and international laws. This includes compliance with all necessary documentation and phytosanitary requirePhellodendron amurense (Amur corktree; fruit shown at left) is currently monitored for its invasive potential in the Arboretum. Spontaneous trees have been removed, and female trees lacking sufficient documentation have been deaccessioned as a means of limiting seed production. However, other individuals of documented origin--some representing unique provenances--remain in the collection because of their high scientific value. NANCY ROSE Living Collections Policy 15 NANCY ROSE Nearly 500 plant genera are common to both North America and eastern Asia. Many representatives of this disjunct group are included in the Arboretum's collection, including two strikingly similar Cornus species, Cornus alternifolia from North America (left) and Cornus controversa from eastern Asia (right). ments during acquisition and distribution activities. All taxa are evaluated for their potential invasiveness, and should invasive or potentially invasive plants be retained for their scientific value, additional management procedures are put into place for containment purposes; they are not distributed for horticultural use. II. SCOPE OF THE LIVING COLLECTIONS The Living Collections are divided into three primary collection categories: Core, Historic, and Miscellaneous Collections; within each are secondary collections. This organization allows priority to be assigned to all extant, as well as potential, accessions within each category, thus guiding collections development, management, and enhancement. It should be noted that none of the primary, or secondary, collections are mutually exclusive and that many accessions fall into multiple categories. A. CORE COLLECTIONS The Core Collections are of highest priority and receive the greatest focus with respect to development, management and enhancement. In general, these collections are intrinsic to the mission of the institution through their research use, and preference is placed on material of documented wild origin. Exceptions to provenance requirements are made only in specific cases when the value is significant enough to warrant accessioning. By and large, these collections are regarded as obligatory. 1. Biogeographic Collections Collections representing the floras of eastern North America and eastern Asia have been an important traditional focus, strongly supporting research related to the floristic relationships between these two regions. In particular, eastern North American-Asian disjunct taxa receive high priority with respect to collections development. 2. NAPCC Collections As part of its commitment to the North American Plant Collections Consortium (NAPCC), the Arboretum maintains and develops collections of botanical taxa RUPERT TAYLOR 16 Arnoldia 66\/1 Interspecific diversity is attained by growing as many species as possible within each of these high-priority genera. To increase intraspecific diversity, we strive to acquire germplasm from multiple provenances of each species so that we may illustrate genetic variation as a function of geographic source. Japanese beech (Fagus crenata) is just one of the species of beech grown as part of the NAPCC collection. MICHAEL DOSMANN within the following genera: Acer, Carya, Fagus, Stewartia, Syringa and Tsuga. Because they serve as national germplasm repositories, development and maintenance maximizes both inter- and intraspecific diversity. 3. Conservation Collections As part of its commitment to the Center for Plant Conservation (CPC), the Arboretum maintains and develops collections of the following species: Amelanchier nantucketensis, Diervilla rivularis, Diervilla sessilifolia, Fothergilla major, Ilex collina, Rhododendron prunifolium, Rhododendron vaseyi, Spiraea virginiana, and Viburnum bracteatum. These species, as well as other taxa of conservation value outside the scope of CPC, are developed and maintained with the goals of preserving as high a level of intraspecific diversity as is practicable. 4. Synoptic Collections Collections of documented wild-origin species that together provide a synoptic representation of the woody flora of the North Temperate Zone are maintained and developed. Emphasis is first placed on generic diversity, and then inter- and intraspecific diversity as is practicable. The goal of a synoptic, or comprehensive, collection is to include the broadest possible representation of the item or group being collected. At the Arboretum this means seeking the greatest breadth across all families that contain woody plants. The Arboretum's synoptic collections cannot contain every woody species, let alone every botanical variety or subspecies, so representative genera and species are selected based on institutional priorities and available space. Living Collections Policy 17 B. HISTORIC COLLECTIONS The Arboretum's early contributions to plant exploration and horticultural improvement are manifested in a number of Historic Collections. In general, these collections are obligatory and maintained, but not actively developed except in cases where authentic material of Arboretum origin can be repatriated or the material is sufficiently unique to warrant accessioning. 1. Arnold Arboretum Accessions Plants collected by early Arboretum staff (e.g., C.S. Sargent, E. H. Wilson, J.G. Jack, J. Rock) may lack sufficient documentation, or be of garden origin. However, because they represent important historical chapters in the development of the institution, they are maintained in the Living Collections. In some cases, these accessions may represent genotypes no longer extant in the wild because of local extinction and thus have high conservation value. 2. Nurseries and Horticulturists Accessions derived from historically significant nurseries, botanical institutions and horticulturists (e.g., H. J. Veitch, T. Meehan, M. Vilmorin) may lack full documentation, but are maintained in the Living Collections. These often represent the initial introductions of species into cultivation and are, in all probability, wild-collected. In some cases, these accessions may represent genotypes no longer extant in the wild because of local extinction and thus have high conservation value. 3. Distinctive Cultivar Collections Early in its development, the Arboretum established diverse collections of garden selections now regarded as cultivars within various plant groups (e.g., dwarf conifers, Malus, Rhododendron, Syringa). Because of their period and oftentimes comprehensive nature, these collections are maintained but not developed. NANCY ROSE In 1885, C. S. Sargent described the goals of the Peters Hill landscape as housing \"a collection for investigation which need not necessarily be permanent.\" Otherwise known as discretionary collections, these have often reflected the research interests of staff scientists. Prior to the substantial Malus collection (shown above), which grew through the work of director Karl Sax and horticulturist Donald Wyman, Peters Hill was home to extensive Crataegus collections--a long-term research project of Sargent. 18 Arnoldia 66\/1 ARCHIVES OF THE ARNOLD ARBORETUM Hydrangea paniculata `Praecox' is an old cultivar with a Latinized epithet. Originally collected in Japan by C.S. Sargent in 1892, this Arnold Arboretum introduction is noted for its precocious floral displays, blooming at least a month before typical plants of the species. MICHAEL DOSMANN 4. Cultivars with names proposed prior to 1953 The Living Collections contain a number of historic cultivars with Latinized names that were proposed in a botanical context prior to 1953. While not developed, these are maintained, particularly when they represent material unique in cultivation. 5. Arnold Arboretum Cultivar Introductions Throughout its history, the Arboretum has selected and introduced a number of clones for ornamental use, many of which were initially regarded as botanical formae but are now recognized as cultivars. Because they arose at the Arboretum, they are maintained and development occurs only to repatriate genotypes lost by the Arboretum. 6. Larz Anderson Bonsai Collection The Larz Anderson Bonsai Collection, while not actively developed, is of high priority within the Arboretum's Living Collection because of its historic and aesthetic value. In addition to housing permanent collections that require high maintenance, The Leventritt Shrub and Vine Garden also displays outstanding ornamentals with exemplary traits. Shown here is accession 178-93-A, Forsythia `Courdijau'. C. MISCELLANEOUS COLLECTIONS In addition to those within the above collection categories, The Living Collections comprise a number of plants grown to achieve display effects, for interpretation, for evaluation, or that may fall outside of traditional scope and not even be accessioned. However, because they play important roles in the Arboretum's research, horticultural and educational work, they are included within the Living Collections. These may be obligatory or discretionary, and development and maintenance decisions are made on a case-by-case basis by the Living Collections Committee. Living Collections Policy 19 SHEILA CONNOR 1. Display Collections Plants of cultivated origin, particularly cultivars selected for unique traits, serve important research and education roles; however their primary value is for display. Examples include ornamentals with exceptional or namental qualities, landscape plants well suited to the New England climate (including those with stress-, insect-, and disease-resistance), as well as those under evaluation. These collections are regarded as discretionary and are developed and maintained as needed, with the acknowledgement that accessions may be deaccessioned when their value no longer meets the appropriate standard. 2. Natural Areas The Arboretum landscape contains several natural areas representative of the New England Flora. Generally, these are maintained through natural regeneration of the present vegetation; however development may occur under certain circum- While they may not contain formally accessioned plants, several natural stances (e.g., restoration fol- areas in the Arboretum (including the North Woods, above) are managed as part of the living collections because of their research potential as lowing major disturbance). well as intrinsic beauty. 3. Spontaneous Flora Spontaneous generation of native, as well as exotic, plants occurs throughout the Arboretum's cultivated landscape. As a matter of course, some of these plants are removed because of their noxious characteristics, some are left in place, while others are accessioned (in particular spontaneous interspecific hybrids or landscape specimens). The forthcoming Policy on the Spontaneous Flora addresses this category more thoroughly. 4. Dana Greenhouse and Nursery Collections A number of plants are cultivated at the Dana Greenhouse and Nursery for experimental, observational, and other programmatic functions outside the scope of production for the accessioned Living Collections. Development and maintenance lies with the primary investigator or other assigned staff member, with the understanding that these may be formally accessioned at a later time. 20 Arnoldia 66\/1 APPENDIX: DEFINITION OF TERMS USED IN THE LIVING COLLECTIONS POLICY An accession is the basic unit of a collection and identified by a unique accession number. By definition it represents a single taxon, from a single source, acquired at one time, and through one means of propagation. An accession may comprise a single plant, or multiple plants, each identified by a letter qualifier following the accession number, or in the case of mass plantings, MASS. Accessioning is the process of adding specimens to the Arboretum's Living Collection and occurs at the time of entry regardless of its stage (e.g., plant, cutting, scion, seed). All accession records are permanent and are not expunged should deaccessioning occur. Acquisition of new accessions may be through field collection, exchange, gift or purchase. All acquisitions must meet specific collections development goals in accordance with the Scope of the Living Collections detailed in this Living Col- The label for accession 638-88-C, Fraxinus tomentosa. lections Policy. A collection is operationally defined as a group of accessions organized by a particular category for curatorial, educational, research, display or other use. A collection need not be physically grouped together, and a single accession may be part of multiple collections. From the perspective of commitment, collections may be discretionary or obligatory. Curation is the process of managing the Living Collections to guarantee its conservation, guide its development, ensure its documentation, and facilitate its enhancement. Deaccessioning is the process of removing a living specimen from the collection, but does not include the removal of any records related to that accession. Deaccessioning decisions are made by the Curator of Living Collections, in consultation with the Living Collections Committee. Development is the process by which the Living Collections undergo change through the acquisition of new accessions and the deaccessioning of accessions no longer needed in accordance with the Scope of the Living Collections detailed in this Living Collections Policy. Discretionary collections can be regarded as temporary or permanent. They meet specific research, display, education or other programmatic needs, but do not necessarily represent collections central to the mission and purpose of the Arboretum. Enhancement is the process of adding value to the Living Collections through documentation, research, and other means. MICHAEL DOSMANN Living Collections Policy 21 Literature Cited Ashton, P. S. 1979. The Director's Report. Arnoldia 39(6): 330369. Dosmann, M. S. 2007. The Arnold Arboretum's living collections: A repository for research. Arnoldia 65(2): 3039. Far rand, B. 1946. Contemplated landscape changes at the Arnold Arboretum. Arnoldia 6(10): 4548. Living Collections Long-Range Planning Committee. 1991. Plants for the Twenty-First Century: A longrange plan. The Arnold Arboretum, Jamaica Plain, MA. Sargent, C. S. 1922. The first fifty years of the Arnold Arboretum. Journal of the Arnold Aboretum 3(3): 127171. Spongberg, S. A. 1979. The collections policy of the Arnold Arboretum: Taxa of infraspecific rank, and cultivars. Arnoldia 39(6): 370376. Wyman, D. 1947. The past year at the Arnold Arboretum. Arnoldia 7(1): 18. Michael Dosmann is Curator of Living Collections at the Arnold Arboretum. Last year a decision was made to deaccession a prominent winterberry holly, Ilex verticillata 22879-F, from its location along Meadow Road across from the Visitor Center. Although it was a noteworthy specimen that consistently produced copious fruits, its large size (nearly 25 feet in diameter) prevented access to one of the original Metasequoia glyptostroboides (524-48-AA). An examination of the records showed that the holly was of unknown origin--not even a nursery source--and that seven other vigorous plants of the accession remained in the collection. MICHAEL DOSMANN The Living Collections comprise all plants formally accessioned, and in a broad sense also contain unaccessioned plants in natural areas, spontaneous flora, and research material. Maintenance, from the standpoint of curating the Living Collections, is the practice of vegetatively repropagating an obligatory accession in order to preserve and perpetuate its genetic lineage. Multiple accessions of the same lineage are genetically identical. Obligatory collections are considered permanent and represent collections central to the mission and purpose of the Arboretum. A taxon (plural, taxa) is a unit of any rank within the taxonomic hierarchy (e.g., family, genus, species, variety, cultivar). SYLVIA WINTER "},{"has_event_date":0,"type":"arnoldia","title":"Ecosystems in Flux: The Lessons of Hemlock Hill","article_sequence":3,"start_page":22,"end_page":28,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25439","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed170bb28.jpg","volume":66,"issue_number":1,"year":2008,"series":null,"season":null,"authors":"Schulhof, Richard","article_content":"Ecosystems in Flux: The Lessons of Hemlock Hill Richard Schulhof en years after the first detection of hemlock woolly adelgid (Adelges tsugae) at the Arnold Arboretum, the hard lessons of biological invasion are written across the face of Hemlock Hill. Large gaps mark the loss of hemlocks, while many survivors, diminished by infestation, stand as relics in growing swaths of successional vegetation. Introduced invasive organisms pose an increasing threat to native biodiversity. As is conspicuously evident on Hemlock Hill, newly arrived pests and pathogens can quickly decimate susceptible native species, creating issues that range from concerns for public access and safety to the long-term management of ecological disturbance. Invasive plant species often follow in the wake of such outbreaks, further disrupting native ecosystems. Responding to invasive species in ways that safeguard people, plants, and the larger environment demands that we more wisely manage the uncertainties of a rapidly changing world. The story of hemlock woolly adelgid (HWA) at the Arnold Arboretum recounts the lessons learned in addressing the rarely predictable, often irreversible consequences of biological invasion. New Invasives: A Steady Parade The scope of the problem is substantial. A 2002 National Academy of Sciences study determined that the USDA inspects roughly 2% of cargo shipments yet intercepts over 53,000 arthropods, pathogens, and plants annually. Although few introduced organisms successfully establish, it is conservatively predicted that 115 non-native insect species and 5 plant pathogens will become naturalized in the United States between 2000 and 2020. Continuing loss of native T biodiversity is recognized as perhaps the greatest long-term consequence of invasive species, which are second only to habitat loss as a primary cause of native species decline in the U.S. Of species on the threatened or endangered list, roughly 50% are at increased risk due to competition or predation from non-native organisms. Some unlisted species, such as the eastern hemlock (Tsuga canadensis), face extirpation or severe reduction over large parts of their range. Each region of the country has its own list of problematic introduced insects and pathogens, with growing public awareness that emerald ash borer (Agrilus planipennis) and Asian longhorned beetle (Anoplophora glabripennis), among others, are dire threats to both cultivated landscapes and native ecosystems. The Home Front It is with some irony that I survey the introduced invasive organisms that today inhabit the Arnold Arboretum. A leader in scientific collecting and importation of plants from east RICHARD SCHULHOF Egg masses in dense, cottony clusters provide a highly visible indication of the progress and intensity of hemlock woolly adelgid infestation. Hemlock Hill 23 RICHARD SCHULHOF Sweet birch (Betula lenta), shown here in golden fall color, is now growing across large areas of Hemlock Hill. As is typical across southern New England, this birch species is a dominant colonizer of the post-hemlock landscape. Asia in the decades before and after 1900, the Arboretum is one of a great many agents that unwittingly introduced species to the North American landscape that later naturalized and wrought destructive impacts. Regardless of our respective \"rap sheets\", the Arboretum and other public gardens now work diligently toward devising management strategies to deal with problematic introduced species. At the Arboretum, developing appropriate responses to invasive species is an ongoing responsibility shared by horticulturists, managers, and administrators. Aggressive incursions of winter moth (Operophtera brumata), garlic mustard (Alliaria petiolata), Japanese knotweed (Polygonum cuspidatum), and other invasives require that we stay abreast of new methods and information, not only to improve the efficacy of our management measures but to do so with ever diminishing environmental impacts. This past fall, the position of Manager of Plant Health was created to coordinate integrated pest management and associated environmental monitoring. Cautionary Tales As we have learned over the years, \"best\" practices are moving targets that shift with increasing knowledge and a changing environment. This can be particularly true in managing recently introduced insects and pathogens whose life cycles, host impacts, modes of spread, and other critical traits may still be relatively unknown. The long-term consequences of various management options are often equally unknown. How we make decisions in the face of uncertainty is of great importance. Confronted with approaching waves of introduced species, what can we learn from previous efforts to manage new invaders? 24 Arnoldia 66\/1 RICHARD SCHULHOF Infested trees on Hemlock Hill in 2003 showing the defoliation and reduction of new growth typical of hemlock woolly adelgid infestation. Hemlock Hill 25 Most recently, the potentially harmful effects of biocontrols--non-indigenous species released to control invasive pests--have received considerable attention. The multicolored Asian lady beetle (Harmonia axyridis), intended to control a range of insect pests, now appears to outcompete and replace some native lady beetle species, while becoming a nuisance in its winter aggregations in homes and buildings. In southern Florida, native Opuntia species are threatened by a South American moth (Cactoblastis cactorum) that had been introduced to control Opuntia naturalizing in the Caribbean. Cases of unforeseen consequence, the nontarget effects of some biocontrols may be remembered as cures worse than the disease. From an earlier period, management response to Dutch elm disease (Ophiostoma ulmi), a public and politically charged effort, targeted its primary vector, the elm bark beetle (Scolytus multistriatus). The American elm's (Ulmus americana) importance as an icon in the cultural landscapes of the Northeast made saving the species a priority for state and municipal agencies, and the resulting massive applications of toxic pesticides contributed to an environmental disaster all too well known today. Past actors on a period stage, decision-makers were undoubtedly influenced by historical biases and limited by critical gaps in knowledge, yet their legacies suggest that response to uncertainty--particularly the consequences of our own actions--merits particular focus today. Managing Hemlock Woolly Adelgid Our ten years of managing hemlock woolly adelgid is a story of decision-making in a rapidly changing informational environment. We began with many uncertainties and traveled a path of pivots and about-faces led by growing knowledge of our own site, analysis of outcomes elsewhere, and key findings from the research community. In 1997 HWA was first detected on the Arboretum's Hemlock Hill, a 22-acre historic natural site whose early public use included frequent visits in the 1840s from Margaret Fuller and other members of the Transcendentalist circle. Prior to infestation, Hemlock Hill was home to over 1,900 eastern hemlocks, some dating to the early 1800s. With its several stands of fully mature hemlock-dominated forest, the Hill had long been appreciated as a place of seemingly wild nature in the midst of the city. The Arboretum was hardly among the first sites to deal with HWA. First detected in Richmond, Virginia in the early 1950s, HWA spread rapidly, decimating hemlock populations in the Mid-Atlantic and coastal Connecticut before reaching Boston. Across much of the range of infestation, the ultimate consequence of HWA was near to complete hemlock mortality within four to twelve years. There were few exceptions. With the prospect of losing one of Boston's most significant natural sites and an integral part of our own history, Arboretum managers addressed challenges of a scope not seen since the 1938 hurricane. The process began with questions. What would be the rate of decline for our hemlocks? How many trees could we protect and at what costs to the larger ecosystem? Could a biocontrol under development save our trees? Although these and other questions would remain unanswered for years, management goals drawn from our organizational mission provided a strong compass for initial decisionmaking. Protecting visitor and staff safety, protecting the larger environment, and preserving a still undetermined number of hemlocks were our key priorities. But where to start? Through the Learning Curve We determined that obtaining reliable, sitespecific information about the spread of the infestation and rates of hemlock decline would be essential to planning an effective management response. Monitoring the health of our hemlocks required mapping the locations and assigning an accession number for each tree. This significant investment was abundantly repaid in data that detailed the progression and severity of the infestation as well as the efficacy of our control efforts; information that continues to inform our decisions. Using assessments of crown health, we evaluated all hemlocks, finding that from 1998 to 2002, the number of trees in poor health increased from 30% to 70%. By 2003, Hemlock Hill was a sickly graygreen color. Data from other sites indicated that we could expect large numbers of hazardous and dead trees within two to three years. 26 Arnoldia 66\/1 That winter we visited forests in Connecticut that had been closed to the public because of the danger presented by hundreds of disintegrating dead hemlocks. Further, we learned that the highly hazardous brittle snags had precluded both salvage operations and efforts to contain rapidly growing populations of invasive plants. Foreseeing similarly grim prospects for Hemlock Hill, we anticipated removing over 1,000 rapidly declining trees within the next two years. Fortunately, that large-scale removal never occurred. The winter of 2004, the coldest in many years, brought several nights with temperatures of -5F or colder, delivering an unexpected reprieve. Although not well documented at the time, HWA is highly vulnerable to extreme cold. Based on surveys at other sites, we estimate that well over 90% of the existing HWA population perished that winter. The following summer, which also brought much needed rain, saw a revitalization of our hemlocks that was a wonder to behold. For once, extreme cold had been a gift, resetting the clock of infestation and allowing more time to find new strategies. Additional changes in approach came with new information from the research community. Publications that elucidated site factors affecting rates of hemlock decline, the relative efficacy of different HWA control methods, and the field performance of highly anticipated biocontrols were part of a burgeoning informational environment that enabled knowledge-based decisions. The Arnold Arboretum was fortunate in that HWA arrived in our vicinity just as many research efforts came to fruition, providing us with essential information that was unavailable to managers of previous infestations. Perhaps our hardest decision thus far concerns the number of hemlocks we attempt to save. The absence of host resistance and limited cultural controls leave us with few management options. Clearly any chemical treatment, even relatively benign horticultural oil, brings concern for the larger environment. At the same time, we are an essential resource for a large urban population that for over 150 years has enjoyed the singular educational and aesthetic experiences of a majestic hemlockdominated forest. Finding balance among stewardship, education, and public service goals, we protect hemlocks that are of sufficient vigor to recover and that grow in conditions that are favorable for treatment and do not present risk of water contamination. HWA is controlled with applications of horticultural oil and, more recently, soil injections of imidacloprid , a treatment now provided to over 40,000 trees at Great Smoky Mountains National Park. We now use this method and pay close attention to ongoing research that monitors for non-target effects and persistence in the environment. Ultimately, it is hoped that these treatments will buy time for the Arboretum's hemlocks until biocontrols or other non-chemical options can offer reliable protection. An ongoing challenge, symptomatic of ecosystem disturbances on a global scale, is the control of non-indigenous plants that often invade when native habitats are affected by introduced organisms. As hemlock mortality continues, canopy gaps become points of colonization for glossy buckthorn (Frangula alnus), Japanese knotweed, and other invasives. Our long-term goal is to promote native hardwood forest where hemlock once grew, and while we actively eliminate invasive vegetation, robust native species, particularly sweet birch (Betula lenta), are rapidly dominating large areas. Adaptive Management Our HWA management strategy continues to evolve, reflecting the iterative learning process needed to develop effective site-specific responses to invasive species. Gathering data that monitor changing conditions as well as the effectiveness of management actions is essential, as is a willingness to completely revise strategies based on new results. Our experience speaks to the value of Adaptive Management, a process developed for the management of complex natural systems characterized by uncertainty. Borrowing from scientific method, it relies on carefully assembled hypotheses, field testing of proposed practices, Hemlock Hill 27 RESEARCH OPPORTUNITIES An unanticipated silver lining was found in emerging research opportunities on Hemlock Hill. The severe consequences of HWA infestation pose compelling questions about the ecological changes associated with decimation of a foundation native species. Beginning a four-year investigation in 2004, the Arboretum collaborated with the Harvard Forest to establish six 15-meter by 15-meter research plots in order to measure the changes occurring when hemlock is abruptly removed from the forest system. We removed hemlocks from four of the plots, with the remaining two left unlogged for use as controls. Measurements established baseline data for soil temperature, available nitrogen, organic soil mass, and understory vegetation. Analysis compared nitrogen cycling, decomposition rates, and regeneration across the six plots. Scheduled to conclude in summer 2008, the study is part of a longerterm Harvard Forest effort to assess ecosystem impacts of HWA in southern New England. A second project examined Chinese hemlock (Tsuga chinensis), a species first grown in North America at the Arnold Arboretum. The research established that Chinese hemlock is cold hardy through at least Zone 6 and is fully resistant to HWA, confirming its suitability as a promising landscape replacement for Tsuga canadensis. P. DEL TREDICI Chinese hemlock (Tsuga chinensis), planted in openings on Hemlock Hill, proved highly resistant to hemlock wooly adelgid. and the monitoring of results to inform next steps and ongoing improvement. It is a model for managing disturbed natural systems that lack both predictability and stability, and for which management outcomes may be determined by variables that are unrecognized or unknowable at the outset--in short, much of the world as we now know it. At the Arboretum, we did not set out to adaptively manage; the approach was born of necessity. But with the appointment of a manager of plant health, we now seek to more fully implement its tenets. Public Awareness The dramatic losses on Hemlock Hill, roughly 30% of the original hemlock population, offer an important local example of a global phenomenon. To build public awareness, the Arboretum now offers school field studies and special tours that explore the fragility of native ecosystems, disturbance caused by invasives, and the complex challenges that result for environmental stewards. As former evergreen forest converts to deciduous woodland, programs will interpret changes in nutrient cycling and species inter- 28 Arnoldia 66\/1 RICHARD SCHULHOF Acknowledgments The author thanks the Arnold Arboretum Hemlock Hill management team for its dedicated efforts: Julie Coop, John DelRosso, Peter Del Tredici, Bob Ervin, Susan Kelley, Alice Kitajima, James Papargiris, and Kyle Port. References Cheah, C., M. Mayer, D. Palmer, T. Scudder, and R. Chianese. 2005. Assessments of biological control of hemlock woolly adelgid with Sasajicymnus tsugae in Connecticut and New Jersey. Proceedings of the Third Symposium on Hemlock Woolly Adelgid in the Eastern United States. USDA Forest Service,Morgantown, West Virginia, pp. 116130. Cowles, R.S., C.A. Cheah, and M.E. Montgomery. 2005. Comparing systemic imidacloprid application methods for controlling HWA. Proceedings of the Third Symposium on Hemlock Woolly Adelgid in the Eastern United States. USDA Forest Service, Morgantown, West Virginia, pp. 169178. Del Tredici, P. and A. Kitajima. 2004. Introduction and cultivation of Chinese hemlock and its resistance to HWA. Journal of Arboriculture. 30:282286. School children examine sweet birch seedlings as part of a new Arnold Arboretum field study investigating the ecological impacts of hemlock woolly adelgid. Dennis, N. 2004. Expanding trade with China creates ecological backlash. Science. 306:968969. Ellison, A.M., M.S. Bank, B.D. Clinton, E.A.Colburn, K. Elliot, C.R. Ford, D.R. Foster, B.D. Kloppel, J.D. Knoepp, G.M. Lovett, J. Mohan, D.A. Orwig, N. L. Rodenhouse, W.V. Sobczak, K.A. Stinson, J. K. Stone, C.M. Swan, J. Thompson, B.Von Holle, and J.R. Webster. 2005. Loss of foundation species: consequences for the structure and dynamics of forested ecosystems. Frontiers in Ecology and the Environment. 9:479486. Holling, C.S. 1978. Adaptive Environmental Assessment and Management. John Wiley & Sons, Chichester, Great Britain. Orwig, D.A., D.R. Foster, and D.L. Mausel. 2002. Landscape patterns of decline in New England due to the introduced hemlock woolly adelgid. Journal of Biogeography. 29:14751487. Pearson, D.E. and R.M. Callaway. 2003. Indirect effects of host-specific biological control agents. Trends in Evolution and Ecology. 18:456461. Pimenthal, D., L. Lach, D. Morrison, and R. Zuniga. 2000. Environmental and economic costs associated with non-indigenous species in the U.S. Bioscience. 50:5365. Simberloff, D. and P. Stiling. 1996. How Risky is Biological Control? Ecology. 77:19651974 Richard Schulhof is deputy director of the Arnold Arboretum. actions. Presentations to the community and feature stories appearing in newspapers and on radio and the web have further disseminated the Hemlock Hill story in Boston and southern New England. Introduced insects and pathogens are here to stay. Looking to the future, warming temperatures will likely enable HWA and other temperature-limited invasives to expand ranges of infestation and more quickly reach lethal densities on host species. The USDA, among other domestic and international agencies, must strengthen efforts to prevent unintended introductions as well as accelerate research programs to better inform management efforts. Institutions such as the Arnold Arboretum, committed to environmental stewardship and with unique expertise, will increasingly contribute to invasive species management. Perhaps more importantly, we can foster awareness, offering our public landscapes as places of witness and learning during a time of remarkable environmental change. "},{"has_event_date":0,"type":"arnoldia","title":"Silver Wins Gold","article_sequence":4,"start_page":29,"end_page":29,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25441","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed1708126.jpg","volume":66,"issue_number":1,"year":2008,"series":null,"season":null,"authors":"Rose, Nancy","article_content":"Silver Wins Gold Nancy Rose T he tallest tree at the Arnold Arboretum isn't a majestic white pine or a venerable beech, it's a silver maple (Acer saccharinum, accession 12560-C). This stately 127-yearold specimen was recently measured at 126 feet (38.5 meters) tall. Its DBH (diameter at breast height) is currently 67 inches (170 centimeters); it takes three people, fingertip-to-fingertip, to encircle the trunk. This tree started its life at the Arboretum in the form of seeds (accessioned under the then-accepted name Acer dasycarpum) received from the nursery of Benjamin M. Watson in Plymouth, Massachusetts on June 1, 1881. Two other silver maples from accession 12560 also lived at the Arboretum for over 100 years, but specimen A was removed in 1982 and specimen B was removed late in 1985 after suffering major damage from the winds of Hurricane Gloria. Acer saccharinum 12560-C displays the typical form of a mature silver maple: a massive trunk that soon divides into multiple upright limbs; thin, pendulous young branches curving up at the tip; and a rounded, spreading crown. The mature bark is characteristically gray-brown, ridged, and scaly. On this tree (and many other old silver maples) the curving bark scales appear to spiral up the massive trunk. The textured bark and impressive girth of Acer saccharinum 12560-C are irresistible to many visitors passing by on Meadow Road; no doubt this is one of the most frequently touched trees in the Arboretum. Acer saccharinum is native to moist woods and river bottoms in much of the eastern half of the United States and a fringe of southeastern Canada. It can grow in drier soils, but may not be as successful or long-lived. Charles S. Sargent noted in Silva of North America, \"On dry and elevated ground...\" silver maple \"...is not handsome...the habit is loose and unattractive....\" No doubt the vigor, longevity, and stature of Acer saccharinum 12560-C is due in part to its ideal growing site in the moist, rich soil of the Arboretum's Meadow area. Silver maple is often considered highly susceptible to storm damage, but Acer saccharinum 12560-C has survived many storms--including the devastating hurricane of 1938--with little damage. Along with other large, old trees at the Arboretum, this specimen is inspected regularly by staff arborists. In 2006, Acer saccharinum 12560-C was tested using radar imaging and wood density borings in addition to visual inspection. The tree proved to be amazingly sound for the most part, but the presence of some decay led to a bit of support work; two cables now connect several of the main vertical limbs, which should help reduce the chance of major limb breakage in high winds. As with most mature trees at the Arboretum, pruning on Acer saccharinum 12560-C is limited to removal of dead wood. To reduce soil compaction (from its many up-close admirers), mulch is spread in a wide swath around the tree and the soil is periodically loosened with a compressed-air tool. Silver maple's popularity as a shade tree has waxed and waned over the decades. Its status as a native plant and its ability to grow quickly in a wide range of soil conditions gave rise to widespread planting in some eras. However, it has just as often been shunned for its irregular trunk habit, susceptibility to storm damage, extensive root system, and prolific seed production. Silver maple is not a good choice for small urban lots or narrow planting strips along streets, but in larger sites such as parks its leafy, shade-casting canopy is an asset. Acer saccharinum 12560-C certainly shows that silver maple can be a beautiful and impressive tree in the right setting. Nancy Rose is editor of Arnoldia. "},{"has_event_date":0,"type":"arnoldia","title":"Index to Volume 65","article_sequence":5,"start_page":29,"end_page":35,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25440","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed170bb6d.jpg","volume":66,"issue_number":1,"year":2008,"series":null,"season":null,"authors":null,"article_content":"Index to Arnoldia Volume 65 Items in boldface refer to illustrations. Ames, Oakes, correspondence with Edgar Anderson 2: 8, 10, 11 Amphibians 1: 3, 10, 1924, 20, 21, 22 -- at Arboretum, chart 1: 24 Anatolia, southern 1: 26, 2831 -- -- hardy cedar from 1: 2635 \"Ancient and Notable Trees of Japan: Then and Now,\" Richard Primack and Tatsuhiro Ohkubo 3: 1021, 1021 Anderson, Dorothy Moore 2: 2, 3 Anderson, Edgar 1: 36 -- -- life of 2: 24 -- -- Balkan expedition 2: 213 -- -- portrait of 2: 4 Anisko, Tomasz 2: 10 Aono, Yasuyuki 2: 1721 Arabidopsis thaliana genome 2: 28, 36 Araucaria cunninghamia, ancient 1: 18, 18 Arboretum, roles of the 2: 2329, 3039; 3: 29 -- best practices for 4: 1617, 27 Arisaema sikokianum 3: 24 Arnold, James 1: 36; 4: 4 Arnold Arboretum, administration 4: 7 -- -- aging accessions 4: 17 -- -- archives 4: 28 -- -- Asia, current research 4: 13, 14 -- -- Beech Path 4: 34 -- -- boxwood accessions and 2: 3, 4, 710, 9, 12, 12 -- -- Bradley Collection of Rosaceous Plants 1: 24; 2: front cover, 40 -- -- Bussey Brook 1: 24; 3: 28, inside back cover; 4: 4 -- -- Bussey Hill 1: 30, 32; 2: 31, 40, inside back cover; 4: 17 -- -- cedars in 1: 26, 30, 3034, 32, 33 -- -- Centre Street 4: 8 -- -- Chinese Path 2: 40, inside back cover -- -- Conifer Path 3: 28, inside back cover -- -- correspondence, online index to 4: 28 -- -- Dana Greenhouses 1: 11; 4: 8, 17 -- -- Deland Endowment 3: 4 -- -- Digital Resources 4: 21, 28, 29 -- -- director's role 4: 23 -- -- ecological change 4: 17 -- -- education 4: 19, 20, 22, 24, 27, 29, 30, 31, 34, inside back cover Abies, and adelgids 3: 2 -- cilicica 1: 31 Academy of Natural Sciences, Philadelphia 3: 7 Acadia National Park, salamanders in 1: 24 Acer 2: 34; 4: 17, 22 -- distribution of 3: 22 -- foliage qualities 3: 22 -- of China 3: 2227 -- acutum 3: 23, 23 -- diabolicum 3: 23 -- griseum 2: front cover, 40, inside back cover -- henryi 3: 24 -- hyrcanum 1: 32 -- maximowiczianum 3: 23, 24 -- miaotaiense 3: 22, 25 -- miyabei 3: 22, 25 -- negundo 3: 22 -- oligocarpum 3: 26 -- palmatum 3: 22, 25 -- -- f. atropurpureum 4: 9 -- pensylvanicum 3: 25 -- platanoides 3: 22 -- rubrum 1: 23; 3: 22 -- saccharum 1: 23; 3: 22 -- schneiderianum 3: 26 -- sinopurpurascens 3: 23 -- wardii 3: 22, 2527, 26 -- -- genetic analysis 3: 25 -- -- in cultivation 3: 25 -- -- Yunnan, collecting in 3: 2627 -- yangjuechi 3: 2224, 2225 -- -- genetic analysis 3: 25 -- -- seeds 3: 22, 23, 24 -- -- Zhejiang, collecting in 3: 2225 A Changing Ecosystem (school curriculum) 4: 30 Adelges tsugae 1: 22; 3: 2, 29 Adelgids, evolution of resistance to 3: 29 Africa, research in 4: 13 Aiello, Anthony S., \"The Quest for the Hardy Cedar-of-Lebanon,\" with Michael S. Dosmann 1: 2635 Ambystomatidae in MA 1: 19 American Association of Botanical Gardens and Arboreta (AABGA), 1983 meeting 2: 35 American Society of Landscape Architects Award 4: 21 -- -- expeditions, archival material on 4: 28 -- -- -- to Asia Minor, 1900s 1: 3132 -- -- -- to Asian hemlock habitats, 200204 3: 29 -- -- -- to Balkans, 1934 2: 213 -- -- -- to China, 1980 2: 40; recent 3: 3; current 4: 1314 -- -- -- to Japan (Wilson's of 1914 revisited) 3: 1020 -- -- Facilities Planning 4: 3, 8 -- -- funding 2: 2329, 38; 3: 4; 4: 20, 23, 26, 29, 32, 33, 34 -- -- governance 4: 20, 2324, 26, 29, 3132, 33, 34 -- -- Head Start partnership 4: 30 -- -- Hemlock Hill 1: 19, 2225; 4: 30 -- -- -- -- salamander study 1: 1925 -- -- hemlocks at 1: 2225; 3: 23, 4, 7, 8, 9, 9; 4: 17, 30 -- -- herbarium collections, digital 4: 28 -- -- history 4: 45, 1920, 21, 22, 23, 34 -- -- -- -- online archives and photographs 4: 28 -- -- horticulture 4: 8, 1617, 1920, 21, 24, 29, 31, 34 -- -- information access 4: 8, 19, 22, 28, 29, 32 -- -- international research, current 4: 1314 -- -- Landscape Institute 4: 20, 24, 29, 33 -- -- Landscape Management Plan 4: 1617, 20 -- -- Leventritt Garden 4: 20, 21, 33 -- -- landscape 2: 23, 26, 34; 3: 28, inside back cover; 4: 79, 1617, 20, 27, 31 -- -- libraries 2: 23, 26; 4: 8, 23, 28, 29 -- -- Lilac Sunday 4: 5 -- -- Linden Path 4: 21 -- -- Living Collections 2: 2329, 3039; 3: 7; 4: 4, 12, 1617, 31 -- -- -- -- accessions added 4: inside front cover -- -- -- -- database 4: 28 -- -- -- -- scientific control 2: 2426, 3138, 31 -- -- -- -- obligatory genera 2: 34; 4: 17, 22 -- -- map, facilities 4: 3 30 Arnoldia 66\/1 -- -- -- online 4: 28 -- -- Meadow 1: 25; 4: 21 -- -- mission and vision 2: 2426, 3338; 4: 34, 12, 1920, 2324, 26, 29, 30, 31 -- -- native tree species plan 1: 23 -- -- neighbors 4: 3, 8, 26, 30 -- -- Oak Collection 2: 23; 4: 2 -- -- Oak Path 4: 31 -- -- Operations Summary 200207 4: 33 -- -- Organizational Chart 4: 32 -- -- paperbark maples in 2: front cover, 40, inside back cover -- -- Peters Hill 2: 30 -- -- photodata 1: 37 -- -- plant health 4: 1617 -- -- public programs 4: 4, 8, 23, 24, 2632, 34 -- -- publishing 1: 3, 10, 32, 36; 2: 24, 25, 26, 33, 36, 38; 4: 19, 28, 4052 -- -- Putnam Fellow 4: 17 -- -- research 1: 29, 1925; 2: 29, 2329; 3039; 4: 39, 1115, 1820, 2324, 26, 29, 31, 34 -- -- -- current international 4: 1314 -- -- -- documenting accessions and 2: 3133 -- -- -- global benefits of new program 4: 15 -- -- Sargent Fellows 2: 29; 4: 1820, 33 -- -- School Programs 4: 30, 30 -- -- seed-collecting 4: 16 -- -- Seed Herbarium Imaging Project (SHIP) 4: 22, 22 -- -- shrubs and vines 4: 20, 21 -- -- staff, list of 4: 3639 -- -- -- published writings of 4: 4052 -- -- Tree of Life project 4: 16, 19 -- -- Visitors 4: 19, 20, 21, 27, 27, 29, 32 -- -- Walter Street 4: 4 -- -- wayfinding systems 4: 21, 27 -- -- Weather Station Data--2007 4: 35 -- -- Weld Hill research facility 2: 25, 26; 4: front and back covers, 312, 1415, 1820 -- -- -- -- -- -- administrative issues 4: 1820, 2324, 26, 29, 31, 34 -- -- -- -- -- -- architect's rendering 4: 11 -- -- -- -- -- -- building description 4: 911 -- -- -- -- -- -- investment 4: 3, 47, 1112, 20, 3034 -- -- -- -- -- -- program 4: 57, 912, 1415, 1820, 31, 34 -- -- -- -- -- -- site planning 4: 3, 39, 67 -- -- wildlife of 1: 19, 2225 \"Arnold Arboretum's Living Collections: A Repository for Research,\" Michael S. Dosmann 2: 3039, 31, 36, 37 Arnoldia 1: 3, 36; 2: 10; 4: 19, 28 -- archives 1: 36; 3: 10; 4: 28 \"Arnoldia Online\" 1: 36, 36 Arundinella 3: 26 Ashton, Peter, research vision for Arboretum 2: 27, 28, 29, 3536; 4: 13 Asia, current research in 4: 13, 14 Asia minor, hardy cedars in 1: 2635 Asian hemlocks 3: 29, 3, 59 -- -- associated flora 3: 7 -- -- distribution map 3: 5 -- -- phylogenetic charts 3: 4, 8 Aspen tree, and polluted land 4: 15 Audubon, John James, Birds of America, illustrations from 1: 23, inside back cover Bagby, Martin 2: 12 Balkans expedition of 1934 2: 213 Balsam woolly adelgid 3: 2 Bamboo 3: 7 -- as impediment to travel 3: 2526 Bank, Mike 1: 24 Barbre, Clarence 2: 12 Barnes, Sheryl 1: 36 Batdorf, Lynn 2: 7, 12 Beech 4: 22, 34 Beijing Institute of Botany 3: 25 Bentham and Hooker classification 2: 34 Betula lenta 1: 23 Biodiversity website, Chinese plants 4: 14, 28 Biofuels, and woody plants 4: 15 Biology, modern approaches 2: 23, 2629, 3037; 3: 34, 78; 4: 1112, 14 Birch, black 1: 23 Birds 1: 3, inside back cover -- and hemlock forests 1: 19, 21, 22, 23 Blandy Research Farm, and boxwood 2: 8, 12 Blue Hill Observatory, long-term temperatures 1: 3, 4 -- -- -- data chart 1: 4 Blue Ridge Mts. 3: 5 Bo tree 3: 14 Boston, City of 4: 3, 3, 4, 8, 20, 30 Botanical Review 2: 30 Botanic gardens, and climate change data 1: 9 -- -- collections crisis 2: 3039 -- -- databases 4: 28 -- -- hemlock research at 3: 29 -- -- leadership 2: 2429 Boufford, David 4: 14 Box elder 3: 22 Boxwood, Balkan 2: 212, 3, 6, 7, 9, 12 -- English 2: 213 -- -- decline 2: 2 -- registrar 2: 7 British Columbia, botanic garden 3: 25 -- -- history of HWA in 3: 4 Broken Arrow Nursery 1: 12 Bussey, Benjamin 4: 4 Bussey Institution 2: 2 Buttercup 3: 26 Buxus sempervirens 2: 213, 3, 6, 7, 9, 12 -- -- Balkan accessions 2: 3, 4, 710, 9, 12, 12 -- -- -- -- herbarium specimen #133 2: 69, 7, 9 -- -- `Inglis' 2: 8 -- -- \"K-Series\" 2: 12, 12 -- -- -- `Agram' 2: 12 -- -- -- `Ipek' 2: 12 -- -- -- `Nish' 2: 12 -- -- -- `Petch' 2: 12 -- -- `Scupi' 2: 8 -- -- suffruticosa `Edgar Anderson' 2: 8 -- -- `Treska Gorge' 2: 8 -- -- `Vardar Valley' 2: 213, 3 -- -- -- -- propagation 2: 67, 8, 9, 10 Cabot Family Charitable Trust 4: 22 California Botanical Garden, Berkeley 2: 35 Camelbeke, Koen 1: 11 Camellia 3: 7 Carbon flows 4: 13 Cardiocrinum lily 3: 26 Cary Arboretum 2: 35 Carya 2: 34; 4: 17, 22 Cedar, Atlas 1: 26 -- Cyprus 1: 26 -- deodar 1: 26 -- of-Lebanon, hardy 1: 2635 -- -- -- Pennsylvania champion 1: 34 -- -- -- in North America 1: 3134 Cedrus, found in wild 1: 26, 30 -- atlantica 1: 26 -- brevifolia 1: 26 -- deodora 1: 26, 29 Index 31 -- libani 1: 2635, 27 -- -- in botanical literature 1: 2635, 26, 27 -- -- ssp. stenocoma 1: 2635, 2830, 3233 -- -- -- propagation 1: 3132, 3334 -- -- -- `Purdue Hardy' 1: 34 Center for Tropical Forest Science (CTFS) 4: 13 Cercidiphyllum 4: 21 -- japonicum, roles in Collection 2: 34 Chemetoff, Alexandre 1: 13 Chemotaxonomic studies 3: 8 Cherry blossom festivals 2: 1416, 1422, 20, 22, back cover -- -- -- charts of flowering times 2: 19, 20, 21 -- Edo-Higan 2: 20 -- Japanese mountain 2: 1618 -- Oshima 2: 20 -- Yoshino 2: 1618 -- -- in Washington D.C. 2: 20, 21 Cherry laurel 2: 4 Chickadee, black-capped 1: 22 Chionanthus virginicus 1: 6 China, arboricultural practices 1: 15, 17, 18 -- expeditions to, 1910 3: 3; 1907 2: 40; 1932 3: 3; 1980 2: 40; recent 3: front covers 1 and 2, 5, 7, 8, 2227, back cover; current 4: 1314 Chloroplast DNA 3: 8, 25 Chollipo Arboretum 3: 8, 9 Cienfuegos Botanical Garden, archives 19461959 4: 28 Climate change 1: 29, 1925; 2: 1422, 37; 3: 4; 4: 13, 15 \"Climate Change and Cherry Tree Blossom Festivals in Japan,\" Richard Primack and Hiroyoshi Higuchi 2: 1422, 1422 Cold-hardy boxwood 2: 213 -- cedar 1: 2635 -- hybrid magnolia 1: 1012 Collections-based research 2: 2329, 3039; 3: 2; 4: 31 -- -- curatorial practices and 2: 3037, 31 Concord [MA] climate change 1: 48 -- -- -- data chart 1: 8 Conservation, plant 4: 21 Cook, Robert E., Director's Report 20032007 4: 335 -- -- -- \"The Future of Research at the Arnold Arboretum\" 2: 2329 Coop, Julie 4: 17 Cornell University 4: 14, 17, 27 Crataegus 2: 34 Cryptomeria fortunei 3: 23 -- japonica 3: 18, 19 Cuba, botanic garden in 4: 28 Cupressus sempervirens 1: 31 Curtis's Botanical Magazine, illustration from 1: inside front cover Cycads in rice straw 1: 16 Cypripedium acaule 1: 5 Daikokuten shrine 3: 20 Daphniphyllum macropodium 3: 24 Database of Asian Plants in Cultivation (DAPC) 2: 35 Databases 2: 35; 4: 28 Davies, Stuart 4: 13 Deer, damage to trees in Japan 3: 18 -- white-tailed 1: 22 Deforestation 1: 23, 30, 31, 32; 2: 6; 3: 2 Del Tredici, Peter 1: 24 -- -- -- \"Magnolia x thompsoniana `Cairn Croft'\" 1: 1012 -- -- -- \"The Paperbark Maple--One Hundred Years Later\" 2: 40 -- -- -- \"`Vardar Valley' Boxwood and Its Balkan Brothers\" 2: 213 -- -- -- photos by 2: covers 1, 2, 3; 3: covers 1, 4 Dendroica fuscus 1: 22 -- virens 1: 22 Desmognathus fuscus 1: 20 Digital imaging of seeds 4: 22 Director's Report 20032007, Robert E. Cook 4: 335 Disjunct flora 2: 27; 4: 17 -- -- and Tsuga 3: 45 Dooley, Major James and Sallie 3: 4 Dosmann, Michael 4: 17 -- -- \"A Golden Afternoon\" 3: 28 -- -- \"The Arnold Arboretum's Living Collections: A Repository for Research\" 2: 3039 -- -- \"The Quest for the Hardy Cedar-of-Lebanon,\" with Anthony S. Aiello 1: 2635 -- -- photo by 3: inside back cover, 4: inside front cover Doyle, Kevin 1: 11, 12 East, Dr. Edward 2: 2 Ecology, tropical forest 4: 13 Ebisudaikoku Pines 3: 20 Edinburgh Botanic Garden 3: 7, 9; 4: 28 Ehwa Woman's University, Seoul 3: 8 Elongate hemlock scale 3: 8 Endangered species 4: 22 Energy independence, and woody plant research 4: 15 Ericaceae 4: 22 Ervin, Robert 4: 16 Europe, hemlock extinction in 3: 4 Eurycea bislineata 1: 20 Evolution, and plant biology 2: 3, 2729, 34; 3: 29, 22; 4: 12 Fagus 2: 34; 4: 17, 22 Fiducela hypoleuca 1: 3 Firs and adelgids 3: 2 Flanagan, Mark 1: 32 Flint, Harrison 1: 34, 37 Floras of Asia 2: 26 -- -- -- and North America 2: 27, 37; 3: 45; 4: 17 Floristics 2: 26, 31 Flowering and climate change 1: 29; 2: 1422 Flycatcher, pied 1: 3 Fordham, Al 4: 22 Forests, Chinese 3: 2226 -- ecology of 1: 1925; 2: 37 -- maple 3: 27 -- hemlock 1: 19, 2225; 3: 28 -- northern hardwood 1: 20, 2225 -- tropical 4: 13 Fortune, Robert, and Pseudolarix 3: 28 Fragrance, in magnolia hybrid 1: 1012 Fringe tree 1: 6 Furlong, John 4: 24 \"Future of Research at the Arnold Arboretum,\" Robert E. Cook 2: 2329, 2326, 29 Gamble, Mary 2: 8, 12 Gansu Province 3: 22 Gao, Lianming 3: 27 Gaoligong, Mt. [China] : 2527 Garden and Forest 1: 10 Gas-exchange experiment 2: 37 Genetics 2: 2728, 31, 35, 37; 3: 3, 4, 79, 22, 23, 25; 4: 12, 14, 19 German language, and plant-collecting 2: 4, 1011, 13 Germplasm repositories 2: 33, 35 Ginkgo, growths on ancient 3: 1617, 1618 -- historic specimens in Japan 3: 12, 1213, 1518, 1518 -- male gametophyte behavior 3: 13 Ginkgo biloba, in China 3: 23 32 Arnoldia 66\/1 Gleason, Herbert Wendell, and flora of Concord 1: 1, 5, 7, 8 Global warming 1: 29, 2122; 2: 1422 \"Golden Afternoon,\" Michael Dosmann 3: 28, inside back cover Golden larch 3: inside back cover Golden-rain tree 2: inside front cover, 37 Greaney-Williams, Ann photo by 4: inside back cover Greenhouse gas 1: 2122 Grimshaw, Scott 4: 31 Groover, Andrew 2: 23, 28 Gyrinophilus porphyriticus 1: 20 Harvard Botanical Museum 2: 26 Harvard Botany Libraries 2: 26 Harvard Dept. of Organismic and Evolutionary Biology (OEB) 4: 5 Harrison, Jim photo by 4: inside back cover Harvard Forest 1: 19, 23; 2: 10 Harvard Graduate School of Design (GSD) 4: 24 Harvard Libraries 4: 23, 28 Harvard Medical School 4: 5 Harvard Online Archival Search Information System (OASIS) 4: 28 Harvard Online Library Information System (HOLLIS) 4: 28 Harvard University 2: 23, 26; 4: 3, 4, 5, 6, 8, 18, 19, 20, 23, 24, 26 Harvard University Herbaria (HUH) 2: 7, 23, 26; 4: 12, 13, 18, 23 -- -- -- specimen 2: 9 Havill, Nathan P. 3: 5 -- -- -- \"The Role of Arboreta in Studying the Evolution of Host Resistance to Hemlock Woolly Adelgid,\" with Michael E. Montgomery 3: 29 Hay, Ida 2: 33 Hebrew SeniorLife 4: 5, 8 Hedera helix var. baltica 2: 4, 8 -- -- -- -- MBG `Bulgaria' 2: 4 -- -- -- -- MBG `Rumania' 2: 4 Heimarck, Heather 4: 24 Helwingia japonica 3: 24 Hemidactylium punctatus 1: 20 Hemlock 4: 22 -- Carolina 3: 2, 5 -- Chinese 3: 23; 2: 37 -- eastern 1: 1925; 3: 2 -- -- HWA nymphs on 3: 2 -- -- replacement species for 2: 37 -- -- soil fauna and 1: 2225 -- mountain 3: 5 -- other pests of 3: 8 -- research in botanic gardens 3: 29 -- Ullung or Korean 3: 8, 9 Hemlock looper caterpillar 3: 8 -- scale, elongate 3: 8 -- woolly adelgid (HWA) 1: 2225; 2: 37; 3: 29, 2; 4: 17, 30 -- -- -- at Arboretum 1: 2425; 3: 23, 7, 8; 4: 17, 30 -- -- -- biological control of 3: 2 -- -- -- genetic variation of 3: 4, chart 4 -- -- -- in evolutionary context 3: 24, 78 -- -- -- in North America 3: 29 Hemlocks, Asian 3: front cover, 29, 3, 5, 6, 7, 9 back cover -- in European fossil record 3: 4 -- taxonomy of 3: 79 Hengduan Mt. region [China] 4: 14 Heptacodium miconioides 2: 35 Herbarium, databases 4: 28 -- specimens 1: 3, 5, 6, 7, 9; 2: 7, 9, 23, 35; 3: 3, 22; 4: 14, 22 Hetman, Jon photo by 4: inside front cover Hickory 4: 22 HighMark Land Design firm 4: 24 Higuchi, Hiroyoshi, \"Climate Change and Cherry Tree Blossom Festivals in Japan,\" with Richard Primack 2: 1422 Himalayas 1: 26; 3: 5 Hohman, Henry 2: 8, 12 Holden Arboretum 4: 16 Honeysuckle, Himalayan 3: 26 Honshu Island 3: 4, 20 Horticulture 2: 27, 35, 36; 4: 8, 1617, 1920, 21, 24, 29, 31, 34 Hosmer, Alfred 1: 6, 7, 8 Howard, Heman 3: 28 Howard, Richard, and research 2: 38 Hubbard Brook Forest, White Mts. [NH] 1: 20 Hubei Province 2: 35; 3: 3, 4 Human Genome Project 4: 12 Hunnewell, Horatio Hollis 3: 28 Hunnewell Pinetum 3: 28 Hurricane of 1938, damage to cedars 1: 32 Hybridization and evolution 2: 3 Hylocichla mustelina 1: 21 Indocalamus, in nature 3: 24 Insects, sucking vs. chewing 3: 8 Invasives 4: 17, 31 Ivy, Baltic 2: 4 Jack, John George, 1905 Japan expedition 3: 15 -- -- -- photos by 3: 15, 16 Japan, ancient trees of 3: 1021, 1021 -- arboricultural practices 1: front cover, 1417, 1417 -- cherry-tree blossom festivals 2: 1422, 1422, back cover -- hemlocks of 3: 29 Jaynes, Dick 1: 12 J. Frank Schmidt Foundation 4: 22 Jiang, Jinhuo 3: 23 John Innes Horticultural Institute [UK] 2: 2, 78 Juniper, Chinese 3: 18, 19 Juniperus chinensis 3: 18, 19 -- foetidissima 1: 31 Kalimantan, research in 4: 13 Kamakura coast pines 3: 13, 1314 -- temple trees 3: 1516 Katsura 2: 34; 4: 21 Keewaydin camp [Ontario] 2: 11 Keteleeria 3: 3 Kingsville Nursery [MD] 2: 8, 12 Kinkakuji Temple, Kyoto 3: 11 Kitaibelia balansae 1: 32 KlingStubbins, Architects 2: 24; 4: 3, 7 -- images by 4: front and back covers, 11 Koelreuteria paniculata 2: inside front cover, 37 Kohl, Paul 2: 12 Koishikawa Botanical Gardens 3: 12 Korean island hemlock 3: 8, 9 Kunming Institute of Botany (KIB) 3: 25, 27 Kyoto, cherries of 2: 1521 -- temple trees of 3: 1012, 1415, 1820 Lady beetle and HWA 3: 23 Lady's slipper, pink 1: 5 Lambir National Park [Malaysia] 4: 13 Land-grant colleges, research role 2: 27; 4: 12, 14 Land reclamation 4: 15 Landscape design profession 4: 24 Larch, golden 3: 28, inside back cover Latin America, research in 4: 13 Leaf emergence and carbon sequestration 4: 15 Lebanon, cedars of 1: 26, 30, 34 Lee, Nam Sook 3: 8 Letea Forest Reserve, on Danube 2: 11 Leycesteria formosa 3: 26 Li, Jianhua 3: inside front cover Index 33 -- -- \"The Search for Two Rare Maples\" 3: 2227 Li, Li 3: 4 Li, Mimi 3: 23, 25 Light-sensing pigment 2: 29 Lilacs 2: 4, 22 Linden, Chinese ancestor in Japan 3: 14, 14 Lindera 3: 26 Linnaean Society 2: 4 Longwood Gardens [PA] 2: 10, 12 Lowell [MA] Cemetery, photodata 1: 2, 24, 8 McCracken Nursery 1: 12 Macedonia, boxwood from 2: 2, 47 Magnolia, first hybrids 1: 10 -- sweetbay 1: 1012 Magnolia `Elizabeth' 4: 25 -- glauca var. major 1: 10 -- tripetala 1: 1012 -- virginiana 1: 10, 11, 12 -- -- `Urbana' 1:10 -- x soulangeana 1: 10 -- x thompsoniana `Cairn Croft' 1: inside front cover, 1012, 1012 -- -- `Olmenhof' 1:11 -- -- zenii 2: 35 \"Magnolia x thompsoniana `Cairn Croft',\" Peter Del Tredici 1: inside front cover, 1012, 1012 Maple, devil 3: 23 -- Japanese 3: 22 -- Miaotai 3: 22 -- Norway 3: 22 -- -- group 3: 23 -- paperbark 2: 40, inside back cover -- pointed-leaf maple 3: 22, 23 -- red 1: 23; 3: 22 -- Schneider 3: 26 -- sheep-horn 3: 2225, 2224 -- stripe bark, American 3: 25 -- sugar 1: 23; 3: 22 -- Tianmu 3: 23 -- Ward's 3: 22, 2527, 26 -- -- in cultivation 3: 25 -- -- in Yunnan 3: 2627 Maples 3: 27; 4: 22 -- eastern North American 3: 22 -- rare Chinese 3: 2227, 2226 -- semi-evergreen 3: 26 -- taxonomic studies of 3: 2227 Mathewson, Brooks, \"Salamanders in a Changing Environment on Hemlock Hill\" 1: 1925 -- -- photo by 1: inside back cover Matthews, Sarah 2: 29; 4: 18, 19 Mayer, Robert G. 1: 19 Maymont Park [VA] 3: 4 Mediterranean region, Cedrus in 1: 26 Meleagris gallopavo 1: 21 Merrill, Elmer Drew 1: 36 Meyer, Henry and Nod 4: 30 Michigan Agricultural College 2: 2 Miller-Rushing, Abraham J., \"Using Photographs to Show the Effects of Climate Change on Flowering Times,\" with Richard Primack, et al. 1: 29 Missouri Botanic Garden 2: 24, 12, 26 Molecular biology 2: 2728; 4: 12, 14 Montgomery, Michael E., \"The Role of Arboreta in Studying the Evolution of Host Resistance to Hemlock Woolly Adelgid,\" with Nathan P. Havill 3: 29 Morris Arboretum 1: 33, 34; 3: 3, 8; 4: 16 Morton Arboretum 4: 16 Moth, winter 4: 17 Mountain habitats 1: 20, 2635; 2: 67; 3: 45, 2324, 2526; 4: 14 Mt. Tianmu Reserve, maples in 3: 2225 Mukunda, Sharda, \"Using Photographs to Show the Effects of Climate Change on Flowering Times,\" with Richard Primack, et al. 1: 29 Muller, Erhart 2: 10, 1011, 13 Muto, master gardener (circa 1911) 3: 4 National Research Fellowship 2: 2 National Science Foundation 2: 27; 4: 16 Necturus maculosus 1: 19 New York Botanic Garden 2: 26, 35 Newt, eastern red-spotted 1: 20, 21 North American Plant Collections Consortium (NAPCC) 2: 34; 4: 17, 22 Nothotsuga 3: 3 Notophthalmus viridescens 1: 20 Nuthatch, red-bellied 1: 22, 23 Nutrient cycling 1: 23 Oak, giant of Nara 3: 18, 20, 20 -- red 1: 23 Oaks, at Arboretum 2: 23; 4: 2, 15 Oakes Ames Orchid Collection 2: 26 Odocoileus virginianus 1: 22 Ohkubo, Tatsuhiro \"Ancient and Notable Trees of Japan: Then and Now,\" with Richard Primack 3: 1021 Olmsted, Frederick Law, landscape principles 2: 34; 4: 78 Omoto, Yukio 2: 1721 Osaka Prefecture University 2: 17, 18 -- flowering cherries of 2: 1721 Pacific northwest, hemlocks in 3: 34 Painter, Jacob and Minshall 1: 34 Panama, research in 4: 13 Panda dens in hemlock 3: 7 \"Paperbark Maple--One Hundred Years Later,\" Peter Del Tredici 2: 40, inside back cover Papua New Guinea, research in 4: 13 Pear, Balkan 2: 4 -- sand 2: 31 Peony 3: 7 People, Places and Design firm 4: 27 Pest management 4: 17 Peters, Wim 1: 11 Phenology 1: 29; 2: 37 Phenotypic plasticity 2: 34 Philadelphia nurseries 1: 34 Photodata 1: 29; 3: 1021; 4: 22, 28 Phragmites, on Danube 2: 11 Phytochrome 2: 29 Pierce, Naomi 4: 13 Pine, Japanese black 3: 13, 13, 18, 19 -- -- white 3: 20, 21 -- white 1: 23 Pinus parviflora 3: 1011, 1112, 20, 21 -- strobus 1: 23 -- thunbergii 3: 1314, 13 Plant biology 2: 23, 2629, 3037; 4: 18 -- -- evolutionary 3: 34, 78; 4: 12 -- hydraulics 2: 29; 4: 18 -- introductions 1: 12; 2: 35; 3: 3; 4: 21 \"Plant Prosthetics: Artifice in Support of Nature,\" Marc Treib 1: front cover, 1318, 1318 Plethodon cinereus 1: 20, back cover Polluted land, and woody plants 4:15 Populus genome 2: 28 Primack, Daniel, \"Using Photographs to Show the Effects of Climate Change on Flowering Times,\" with Richard Primack, et al. 1: 29 Primack, Richard, \"Ancient and Notable Trees of Japan: Then and Now\" 3: 1021 -- -- \"Climate Change and Cherry Tree Blossom Festivals in Japan,\" with Hiroyoshi Higuchi 2: 1422 -- -- \"Using Photographs to Show the Effects of Climate Change on Flowering Times,\" with Abraham J. Miller-Rushing et al. 1: 29 34 Arnoldia 66\/1 Primrose 3: 7 Propagation research 4: 22 Prunus jamasakura 2: 16, 20 -- -- flowering and temperatures 2: 1422 -- laurocerasus var. shipkaiensis 2: 4, 8 -- pendula f. ascendens 2: 20 -- serrulata 2: 16 -- -- var. speciosa 2: 20 -- -- var. spontanea 2: 16 -- x yedoensis 2: 1921 Pseudolarix 3: 3 -- amabilis 3: 28, inside back cover -- -- cone 3: 1 -- -- cultivation at Arboretum 3: 28 -- -- tall `Nana' cv. 3: 28 -- -- wild-collected 3: 28 Purdue University 1: 34 Pyrus elaeagrifolia 2: 4 -- pyrifolia 2: 31 Quarry Hill Botanic Garden 2: 35 Quercus 4: 2 -- gilva 3: 18, 20 -- rubra 1: 23 \"Quest for the Hardy Cedar-of-Lebanon,\" Michael S. Dosmann and Anthony S. Aiello 1: 2635, 2630, 3233 Radcliffe Seminars in Landscape Design 4: 20, 24 Ranunculus 3: 26 Red eft, toxic defense and moisture 1: 20, 20 Reed|Hilderbrand firm 4: 21 -- -- site plan of Weld Hill by 4: 67 References, digital or online 4: 14, 22, 28 Reforestation efforts 1: 34; 4: 13 Research, at Arboretum 2: 2339; 4: 39, 1115, 1820, 2324, 31, 34 Rhododendron 3: 7, 26 Rice straw, and Japanese tree culture 1: 16, 1617; 3: 1516, 16 Richmond [VA], and introduction of HWA to North America 3: 4 Robin, American 1: 21 Rock, Joseph: 3: 2, 3; digital archives 4: 28 \"Role of Arboreta in Studying the Evolution of Host Resistance to Hemlock Woolly Adelgid,\" Nathan P. Havill and Michael E. Montgomery 3: 29, 29 Roll Barresi firm 4: 27 Romania (Rumania) 2: 2, 78, 10, 11 Royal Botanic Garden, Edinburgh 3: 7, 9; 4: 28 Rubus, invasive 4: 31 -- lasiostylus var. hubeinsis 2: 35 Salamander, blue-spotted 1: 19 -- dusky 1: 20, 24 -- eastern red-back 1: 1922, 2325, back cover -- -- abundance at Arboretum, chart 1: 24 -- -- forms of 1: 19, 20, 2022, 24 -- four-toed 1: 20, 25 -- Jefferson 1: 19 -- mole type 1: 19 -- mudpuppy 1: 19 -- northern spring 1: 20, 25 -- -- two-lined 1: 20 -- species in New England 1: 1925 -- vernal activity 1: 1920 \"Salamanders in a Changing Environment on Hemlock Hill,\" Brooks Mathewson 1: 1925, 1924 Samurai and trees 3: 11, 12, 16 Sargent, Charles Sprague 1: 10, 31, 36; 2: 3334, 40; 4: 4, 12, 13, 19, 24, 31, 34 Sargent Fellows 2: 29; 4: 1820, 33 Scale, elongate hemlock 3: 8 Schneider, Steve 4: 16 Schulhof, Richard 1: 24; 4: 32 -- -- \"Landscape Institute\" 4: 24 -- -- \"Linden Path and the Leventritt Garden\" 4: 21 -- -- \"Management Initiatives for Collections and Landscape\" 4: 1617 -- -- \"School Programs\" 4: 30 -- -- \"Seed Herbarium Imaging Project\" 4: 22 -- -- photos by 4: inside front cover, inside back cover Scientific research, prospects for 2: 2339; 4: 39, 1115, 1820, 2324, 31, 34 Scymus sinuanodulus 3: 2 \"Search for Two Rare Maples,\" Jianhua Li 3: 2227, 2226 seed morphology, imaging 4: 22 Shaanxi Province 3: 4 Shaw, Ted 1: 34 Shigeta, Mayumi, photograph by 2: back cover Shinagawa, Fujiro 2: 22, 22 \"Ship Pine\" of Kyoto 3: 1012, 10, 11 Shiyake, Shigehiko 3: 4 Shogun's Ginkgo, Tokyo 3: 1213, 12 Sichuan Province 3: front cover, 4, 5, back cover; 4: 14 Siehe, Walter 1: 31, 32 Silva, online archives 4: 28 Sino-American Botanical Expedition, 1980 2: 35; 3: 2 Sitta canadensis 1: 22 Snakes, garter 1: 21, 24 Soil fauna 1: 1925 Sorbus umbellata 1: 32 -- yuana 2: 35 South Korean hemlock 3: 8, 9 Sparks, Tim 1: 3 Spicebush 3: 26 Stachyurus chinensis 3: 24 Staking methods 1: front cover, 1318, 1318; 3: 1011, 1011 -- contemporary European 1: 13, 13 Stanley Smith Horticultural Trust 4: 22 Staphylea holocarpa var. rosea 2: 36 Stewartia 2: 32, 34; 4: 17, 22 -- phylogenetic chart 2: 32 -- ovata var. grandiflora, seeds 4: 22 -- sinensis 3: 23, 24 Sugi tree 3: 18, 19 Syria, cedars in 1: 26, 30, 32 Syringa 2: 34; 4: 17, 22 `Purple Haze' 4: 17 -- vulgaris 2: 34 Taiwan 3: 4 Takakannon Gonshoji Temple, Kyoto 3: 1415 Takao, Mt. [Japan] 2: back cover Taurus Mts. [Turkey] 1: 26, 28, 2832, 34 Taxa 2: 29, 32, 3637 -- rare 4: 22 Taxonomic studies 2: 3437; 3: 45, 79 Taxus baccata 1: 31; 2: 4, 8 Terpenoids in hemlocks 3: 8 Thamnophis sirtalis 1: 21 Thoreau, Henry David, wildflowers mentioned by 1: 5, 7 Thrush, hermit 1: 21 Thuja plicata 4: 19 Tianmu Reserve [China] 3: 2225 Tibet, research in 4: 14, 28 Tilia 4: 21 -- miqueliana 3: 14, 14 Index 35 Toad, American 1: 24 Tokyo, flowering cherries in 2: 14, 15, 16, 20, 22 -- landmark trees of 3: 12, 12, 1618, 1618 -- botanic gardens 3: 12 Tonkin, John 1: 34 Topyedik, Mt. [Turkey] 1: 28 Towada volcanic caldera 3: 20 Trees, biology of 2: 2629, 3334; 4: 12 -- ancient of Japan 3: 1021 -- evolution and biology -- giant Asian 3: 12, 1620, 23 -- rare Chinese 3: 2227, 28 -- sacred to Buddhists 3: 14, 16 -- veneration of, in Asia 1: 17; 2: 1418; 3: 1021 -- war-damaged 3: 12, 13, 18 Tree-support practices 1: front cover, 1318, 1318; 3: 1011, 1011 Treib, Marc, \"Plant Prosthetics: Artifice in Support of Nature\" 1: 1318 -- -- photo by 1: front cover Trends in Plant Science 2: 23 Treska Gorge [Macedonia] 2: 59 -- -- map 1937 2: 5 Tropical forests 4: 13 Tsuga 4: 17, 22 -- adelgid resistance and evolution 3: 29 -- biogeography 3: 45, chart 4 -- distribution 3: 5, map 5 -- elevation and 3: 5, 8 -- fossil record 3: 4 -- hybrids 3: 8 -- new species 3: 9, 9 -- other pests of 3: 8 -- panda dens in 3: 7 -- phylogenetic chart 3: 8 -- canadensis 3: 2, 45, 7, 8 -- -- `Albo-spica' 3: 8 -- -- cultivars 3: 8 -- -- `Snowflake' 3: 8 -- -- woolly adelgid and 3: 2, 7, 8 -- caroliniana 3: 2, 5, 8 -- chinensis 2: 24, 37; 3: 3, 5, 7, 8 -- -- var. tchekiangensis 3: 6 -- diversifolia 3: 5, 7, 8, 9 -- dumosa 3: front to back covers, 3, 5, 7, 7, 8 -- formosana 3: 8 -- forrestii 3: 5, 5 -- heterophylla 3: 5 -- mertensiana 3: 5 -- sieboldii 3: 5, 6, 8, 9 Tsurugaoka Hachimangu Shrine 3: 1516, 1516 Tsurugi, Mt. [Japan] 3: 6 Turdus migratorius 1: 21 Turkey (Anatolia), cedars from 1: 2635 Turkey, wild 1: 21 Tyler Arboretum 1: 34 Ullung Island hemlock 3: 8, 9 Understory vegetation 1: 2223, 31, 32; 3: 57, 20, 24, 26; 4: 31 University of British Columbia, Botanic Garden 3: 25 University of Illinois 1: 10 University of Memphis 3: 7 University of Pennsylvania 4: 16 University of Toyko, garden 3: 12 University of Washington, Seattle 2: 12 University of Wisconsin 3: 2 Urban forest 1: 23 -- heat 1: 4; 2: 15, 17, 1922 USDA 2: 2, 27, 35; 3: 5; 4: 14 USDA Forest Service 3: 5 US National Arboretum 2: 7, 8, 12; 3: 3, 7, 8; 4: 16 US National Collection of Insects 3: 3 \"Using Photographs to Show the Effects of Climate Change on Flowering Times,\" Richard B. Primack, Abraham J. Miller-Rushing, Daniel Primack, and Sharda Mukanda 1: 29, 2, 4, 5, 6, 7, 8 Van Meulder, Jef 1: 11 Vardar Valley [Yugoslavia] 2: 47 -- -- map, 1937 2: 5 \"`Vardar Valley' Boxwood and Its Balkan Brothers,\" Peter Del Tredici 2: 213, 37, 912 Vegetative succession 3: 20 Veitch and Sons nursery 3: 28 Vireo, solitary 1: 22 -- yellow-throated 1: inside back cover Vireo solitarius 1: 22 Warbler, Blackburnian 1: 22, 23 -- black-throated green 1: 22, 23 Ward, Frank Kingdon, and discovery of Acer wardii 3: 25 Warsowe, Julie 4: 27 Washington Park Arboretum, Seattle 2: 12 Watershed dynamics, and climate change 4: 13 Weather data 1: 29; 2: 1418; 4: 35 Weather Station at Arboretum--2007 summary 4: 35 Weld Hill Research Facility, proposed for Arboretum 4: front and back cover, 312, 3, 6, 11, 1415, 1820, 33, 34 Westwood [MA] origin of magnolia hybrid 1: 1112 Wharton, Peter 3: 25 White, Orland 2: 8 Wilson, E.H. 1: 32, 36; 2: 31, 37, 40; 3: 2, 3 -- -- -- Japanese trees revisited 3: 1021 -- -- -- photographs by 3: 10, 1221 Windsor Great Park, research sponsored by 1: 32 Winter moth 4: 17 Wood, structure of 4: 15 Woodiness, evolution of trait 2: 2829 World War II, remembered in Asia 3: 13, 18, 2627 Wyman, Donald, Arnoldia and 1: 32, 36 -- -- boxwood and 2: 8, 10 -- -- cedar and 1: 32 Xiaohua, Jin 3: 2526 -- -- photo by 3: inside front cover Yale Institute for Biospheric Studies 3: 4 Yale University 3: 3 Yew, European 2: 4 Yu, Guoyue 3: 4 Yugoslav Forest Service 2: 12 Yugoslavia 2: 2, 410, 12 Yunnan Province 3: inside front cover, 4, 5 -- maples in 3: 2526 Zelkova serrata 4: inside front cover Zenpukuji Temple, Tokyo 3: 16, 17, 18 Zhejiang Province 3: 2225, 28 Zhejiang University 3: 25 Zhou, Jianhua 3: 4 Zhao, Mingshui 3: 2324, 24 Zuk, Judy 2: 33 Zwieniecki, Maciej 2: 29; 4: 18 "},{"arnoldia_cover":true,"has_event_date":0,"type":"arnoldia","url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=23409","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9dcabd25e856e.jpg","title":"2008-66-1","volume":66,"issue_number":1,"year":2008,"series":null,"season":null},{"has_event_date":0,"type":"arnoldia","title":"Director's Report: 2003-2008","article_sequence":1,"start_page":1,"end_page":53,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25437","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed170b36b.jpg","volume":65,"issue_number":4,"year":2008,"series":null,"season":null,"authors":"Schulhof, Richard; Cook, Robert E.","article_content":"THE ARNOLD ARBORETUM OF HARVARD UNIVERSITY DIRECTOR'S REPORT: 2003?2007 Robert E. Cook, Director ARNOLDIA ? VOLUME 65 ? NUMBER 4 Arnoldia (ISSN 004-2633; USPS 866-100) is published quarterly by the Arnold Arboretum of Harvard University. Periodicals postage paid at Boston, Massachusetts. Copyright ? 2008. The President and Fellows of Harvard College. The Arnold Arboretum of Harvard University 125 Arborway, Boston, Massachusetts 02130 FRONT COVER: Weld Hill research facility, design sketch of Centre Street view (detail); KlingStubbins. BACK COVER: Model of Weld Hill research facility by GPI Models; photographs by Desroches Photography. Top main entrance and laboratory wing on the north side of the building; Bottom courtyard and greenhouses on the south side of the building. Quercus (oak) collection by Jon Hetman Introduction E arly this spring, the Arnold Arboretum began construction of a new research and administration building at Weld Hill, a fourteen-acre parcel of land adjacent to the grounds of the Arboretum (see Figure 1). It will be the first major building added in nearly half a century. The Weld Hill facility, as we are calling it for now, will have nearly 44,000 square feet of floor area and cost approximately $42,000,000. Its greenhouses, growth chambers, and modern laboratories will provide state-of-the-art facilities for plant research. The construction of the building marks a major milestone in the history of the Arboretum and a reaffirmation of our mission as a research institution at Harvard University. In this Director's Report, I will focus on a physical description of the building and its location, the decisions that led to its construction, and the implications of its Figure 1. Map of the Arnold Arboretum, showing Harvard-owned land, City of Boston-owned land leased to Harvard, and the proposed facility additions (see Facilities Planning, page 8), by KlingStubbins. 4 operation on the future programs of the Arboretum. At the end of the report, I will return to the critical role that all the friends of the Arboretum have had in reaching this milestone. A Building in its Landscape A s is well known, most of the land of the Arboretum is owned by the City of Boston and open to the public as part of the Boston park system; however it did not begin this way. Through the generosity of James Arnold, and on land donated by Benjamin Bussey, Harvard University created the Arboretum as a private research department dedicated to the study of trees. Within a decade the first director, Charles Sprague Sargent, had developed a unique partnership with the City that captured his vision for the institution. It was to become both a non-public research station and a public museum. A decade later, in 1882, the Arboretum's land was given to the City to be operated as a park open to the public for education and enjoyment. At the same time, the land was leased back to the University, for a fee of one dollar a year and a renewable term of 1,000 years, to allow faculty and students to conduct research on the biology of trees. This partnership between Harvard and the City has successfully endured up to the present day. The Weld Hill building is simultaneously an affirmation of the partnership and the embodiment of Sargent's original vision. Several considerations determined the choice of location for the facility. First, it was highly desirable that a new facility not subtract from the parkland available to the public; a location on the grounds would have had this effect. At the same time, however, researchers would highly value close proximity to the collection of trees upon which their research is conducted. Finally, in consideration of both cost and formal control of the facility's operation and construction, we preferred to site the building on land already owned outright by the University, close to but not within the historic public park. In 1922 Charles Sargent, in his fiftieth year as director, purchased from a neighbor a fourteen-acre parcel of land lying across Walter Street and adjacent to the southwest boundary of the Arboretum (see Figure 2). The parcel displays significant topographic variation, from the drainage swale feeding Bussey Brook to a large hilltop that reaches 172 feet above sea level. Over the subsequent eighty years, it was variously used for nurseries, tree plantations, and the planting of some specimen trees. By 2002 Weld Hill's pastures and woodlots were no longer home to any significant collections and horticultural care had been reduced to an annual mowing of its meadows. To the north of Weld Hill, a major teaching and research hospital for the elderly, called Hebrew SeniorLife, had become an important partner of the Harvard Medical School. Following completion of our strategic plan in 2002 (see Director's Report 1999?2002), we decided that this parcel of land would become the home for a laboratory facility representing a major expansion of our capacity to conduct research. The functional requirements were developed collaboratively with botanical faculty members in Harvard's Department of Organismic and Evolutionary Biology (OEB), with which the Arboretum has worked for more than a century. These requirements grew out of the Arboretum's commitment, described in the 2002 Director's Report, to make a strategic investment in scientific research.1 Our goal for 1 R. E. Cook. \"The Future of Research at the Arnold Arboretum,\" Arnoldia 65(2): 23?39 (2007). Lilac Sunday visitors by Eric Roth. 6 Figure 2. Site plan with topographic detail and building footprint for the Weld Hill facility, by Reed|Hilderbrand. this investment was to match other major research units at Harvard University in providing the physical and financial capacity to support researchers of the highest quality. Nothing pleases high-quality researchers more than first-class research facilities. For plant scientists, this means the finest growing facilities (controlled climate greenhouses; growth chambers; nurseries and experimental gardens). And, if at all possible, it also means access to a broadly diverse collection of living 7 tree species of great maturity and known provenance from all over the world. The scientific program of the building must support a sufficient number of scientists to form an intellectual community that interacts and collaborates closely. Toward that end we specified enough laboratory space to house eight senior researchers or faculty members, along with their post-doctoral trainees, their graduate students, their laboratory technicians, assorted undergraduates just getting started, and visiting scientists. We estimated the total to be about 40 full-time researchers. In addition, as part of a larger facilities plan to address other needs at the Arboretum (see Facilities Planning, page 8), the new building would also accommodate the relocation of the Arboretum's administrative offices, presently housed in the Hunnewell Building. These offices include the director's and those of such functions as finance, facilities management, personnel, and research administration. The space required for these purposes, together with the mechanical systems to heat and cool the facility, comes to approximately 43,500 square feet. Taking into account site conditions (topography, wetlands, utility easements, rock) and the necessary road and parking infrastructure, we envisioned that the facility would require less than a quarter of the entire fourteen acres of the Weld Hill site. In collaboration with the architectural firm of Stubbins Associates (now KlingStubbins, Inc.), located in Cambridge, MA, we defined a set of principles for siting the building that would be consistent with the pastoral nature of the Weld Hill land and with the Olmsted FAC I LITI E S PL AN N I N G I n 2004 the Arboretum developed a comprehensive facilities plan. At its center was the creation of the research facility that is described in this director's report. But the plan also projected the construction of facilities that would address two other needs. First, our primary maintenance facility, which houses grounds equipment adjacent to the Hunnewell Building, has reached the end of its useful life. In the plan we proposed a new facility that would be more centrally located in the vicinity of the Dana Greenhouse, thereby separating the traffic in tractors and trucks from the movement of pedestrians around the Hunnewell Building Visitor Center. Second, we wanted to expand our service to the public adding a new wing to the Hunnewell Building that would house our educational programs and visitor services and provide greater access to the horticultural library. Thus the larger facilities plan envisioned three centers (see Figure 1). A new research and administration complex would be located at Weld Hill. The horticultural support facility would include a new maintenance building combined with the Dana Greenhouse. Finally, the Hunnewell Building, with a newly constructed public wing, would be the \"public museum,\" dedicated to education and service. However, late in 2005, after developing schematic designs for the maintenance facility at the Dana Greenhouse, we suspended any further work on it and, consequently, on any expansion of public programs at the Hunnewell Building as well. Our plan called for the removal of an 1820 farmhouse, in dilapidated condition and of no further use to the Arboretum, because its location precluded the construction of a maintenance facility on land owned outright by Harvard University rather than on land under lease from the City and historically part of the public realm. A small group of neighbors objected to our proposed removal for reasons of historic preservation. We made the strategic decision to focus our immediate energies on the construction of the Weld Hill facility. design of the adjacent Arboretum landscape. These principles are: 1) to locate buildings on edges and in corners; 2) to concentrate buildings and circulation infrastructure near other buildings and infrastructure; and, 3) to reserve the higher elevations for pedestrians and use the lower, level land for nurseries. These principles, combined with the concern expressed by neighbors that they not \"see\" the building, led to its current planned location in the parcel's northwest corner on the lower slope of Weld Hill (see Figure 2). Here the building will lie close to the existing hospital facility (Hebrew SeniorLife) and be terraced into the lower hillside such that the mass and height of the hill will screen the building from its neighbors to the south. The service road will enter the site from Centre Street and curve across and around the rising topography to meet the rear of the building at its highest floor, adjacent to the greenhouses, before descending to an exit on Centre Street. This siting effectively leaves the hill itself open for pedestrian enjoyment and reserves the eastern half of the parcel for woodlands, pastures, and gardens. 9 Thus the overall landscape will be characterized by its four quadrants. The northwest quadrant will hold the building and its circulation infrastructure; the pastures and distant views of the hilltop will continue to occupy the southwest quadrant; mature woodlands will remain throughout the northeast quadrant; and the southeast quadrant will display terraced nursery beds. A Home for Research T he facility itself will be a large, three-story central building with Acer palmatum f. atropurpureum (AA 22717-A), two wings to the east and west photo by Jim Harrison. (see Figure 3). The basement floor of the central building will consist of a large room for mechanical equipment adjacent to the main entrance and the stairway leading to the administrative offices, computational facilities, and a large auditorium for meetings and lectures on the first and second floors. The first floor of the east wing will have a large, open laboratory conducive to collaborative research, with individual work stations on the north side and specialized equipment rooms tucked into the hillside on the south. Above this, on the second floor, a series of offices will house senior researchers and faculty. The west wing will be devoted to growing plants. On the first floor, several large rooms will be equipped with reach-in and walk-in growth chambers, as well as related laboratory support spaces. The second floor will have a large head house and loading dock to support twelve adjacent greenhouse modules. My simple description of these spaces with their highly specialized equipment and technical capacity does not evoke the 10 ambiance of the building--the way it will look and feel as it sits in its landscape. I wanted the building's occupants to feel very much at home despite the essentially institutional nature of their work. This is a challenge for a fairly large facility that one might initially envision as a steel and glass box like most laboratories being built today. The configuration of the interior spaces will be largely dictated by functional needs; any domestic qualities must evolve from the ingenuity of their occupants. Nevertheless, we have tried to communicate in the design of the building's exterior--its appearance and surface qualities--a sense of arriving at home as one approaches the front door. The fa?ade of the building will be constructed of basic, traditional, residential materials: stone, brick, and wood. The lowest level of the central building will be clad in light Kasota stone of a lemon hue. This stone base will rise to meet horizontal shiplap cedar siding covering the first and second floors, both of which will display horizontal bands of vertical windows (see Figure 3). The base of each wing will be articulated in brick that rises to meet cedar-clad walls and rows of windows. At the end of each wing and at the junctures with the central building will be large vertical brick \"chimneys\" to house mechanical ducts and other equipment. Most important, the roofs of the entire building will slope gently down to create substantial overhangs. They will be shingled with a dark synthetic slate, and the gutters, downspouts, and building accents will be made of copper. The overall appearance of the building will recall the early prairie houses of Frank Lloyd Wright, particularly Taliesin East, his first studio and home in Wisconsin. With its central mass flanked by two recessed wings that recede into the slope of the hill; its strong horizontal lines defined by roof, windows, and base; and its cladding of natural materials, the building will be fully integrated with and defer to the surrounding pastoral landscape. The immediacy of the landscape will be reinforced on the south side of the central building by a courtyard sanctuary that visually connects the first-floor reception area to the summit of the hill (see Figure 3). The design and engineering of the facility and its site will fully incorporate current principles of sustainable architecture. The roof shingles, for instance, will be made of recycled tires, and other materials Figure 3. Architectural rendering of the north (top) and south views of the Weld Hill facility, by KlingStubbins. throughout the building have been chosen to minimize their negative impact on the environment. Water flow from rain falling on the building and land will be managed to minimize any change to the hydrological conditions that prevailed prior to construction. Rooms will have ceiling fans and all windows will be operable to maximize natural circulation and encourage energy conservation. Finally, and most significantly, the building will be heated and cooled by a geothermal exchange system buried deep below the ground. This system--essentially a network of connected pipes buried in deep bore holes--will work like a very large radiator drawing heat from the earth in the winter and returning it in summer. There will be no basement furnaces or rooftop air conditioners. Building Research Capacity B ecause of the challenges presented by the building's hillside location and its environmentally sensitive architecture, the Weld Hill facility is relatively expensive on a cost-persquare-foot basis. Can such an investment in research at the Arnold Arboretum be justified? I believe the answer is yes. Elsewhere I have written extensively on the revolution sweeping the botanical sciences and the critical 12 importance of the Arboretum's participation in this revolution.2 Let me briefly review the reasons for our investment at this time. In 2001 a landmark goal was reached in biomedical science when the Human Genome Project completed the sequencing of all human genes. Out of this work and related efforts with other species' genomes have come whole new approaches to basic research that are fundamentally transforming our understanding of biological organisms and their evolution. A new field of science, evolutionary developmental biology, has emerged that embodies many of the elements of this revolution. It seeks to understand how a sequence of genes inherited from two parents leads to the creation of a new organism and how this system of reproduction evolved over time. The methods of research in this new field, while drawing deeply on the sciences of molecular biology and biochemistry, have also given renewed importance to comparative biology, which analyzes the differences between species and studies their evolutionary history in order to bring insight to discoveries in developmental biology and genetics. All of these new approaches and their highly technical methodologies apply to the biology of plants as well as that of animals. This has three major implications for the Arnold Arboretum. First, if the Arboretum is to maintain its scientific reputation for the long-term future, it must invest in the capacity to conduct this kind of research. The original trustees who created the Arboretum at Harvard University and the Arboretum's first director, Charles Sprague Sargent, would have surely agreed. Second, with its collections and endowment, the Arboretum has an exceptional opportunity to advance our understanding of plant biology, particularly that of trees. The herbarium and living collections, in addition to our incredible library collections, can only become more valuable as tools to further our understanding of comparative biology and the evolutionary history of botanical diversity. And, as noted earlier, the Arboretum needs research facilities of the highest caliber to preserve and enhance our future scientific reputation. Should the Arboretum conduct applied research with more targeted goals typical at agricultural colleges and land-grant universities? I believe the answer is no, because the highest-quality research, 2 Ibid. C U R R E N T I N T E R N AT I O N A L R E S E A R C H A lthough the Arboretum is investing in a major research expansion at Weld Hill, it continues its international research in Asia through projects in southwest China, Kalimantan (Indonesian Borneo), and Papua New Guinea. Also of critical importance is our growing collaboration with the Smithsonian Institution. It was Charles Sprague Sargent, the Arboretum's first director, who first recommended an expansion of the Arboretum's mission to the tropical forests of Asia.1 This tradition reached its apex in the spring of 2007 when Peter Ashton, Bullard Professor Emeritus and director of the Arboretum from 1979?1987, was awarded the Japan Prize for his lifetime devotion to research on the biology of Asian tropical forests (see Director's Report 1997?1999 for a full description of this work). In 1983 Peter initiated a collaboration between the Smithsonian Institution's Tropical Research Institute (STRI) and the Arnold Arboretum which has since evolved into the Center for Tropical Forest Science (CTFS), a network of international partners around the world. This program supports long-term tropical forest research through a set of permanent, large-scale plots established in forests that differ in climatic conditions, soil types, and disturbance regimes. These research plots, located at twenty sites in fifteen different tropical countries in Asia, Africa, and Latin America, are united in maintaining a standardized monitoring methodology involving a complete census every five years of all trees larger than one centimeter in diameter, with each individual being mapped, tagged, measured, and identified to species. The first plot was estab1 lished on Barro Colorado Island in Panama in 1980. In recent years the number of permanent plots has been expanding through support from the Arnold Arboretum, particularly in tropical Asia. Recently the Center received grants to create a global earth observatory system for research on forest dynamics in response to climate change. This system will be based on the already existing permanent tropical forest plots and expanded to include new plots in temperate regions of the world. The grants will support measurement and monitoring of carbon flows and watershed dynamics in forests that are experiencing the impact of changes CTFS director Stuart Davies and in global climate. Harvard biology professor Naomi The Center also Pierce at the CTFS tropical forest plot maintains programs at Lambir National Park, Malaysia, photo by Christian Ziegler. of field training, research grant support, and applied research into sustainable policies for the management and restoration of tropical forests. In June of 2007 the headquarters of CTFS, and its director, Dr. Stuart Davies, relocated to the Harvard University Herbaria, with the Arnold Arboretum assuming full support for the twelve permanent plots in Asia. The CTFS program will continue to maintain the core of its operations at STRI in Panama. C. S. Sargent. \"The First Fifty Years of the Arnold Arboretum,\" Journal of the Arnold Arboretum 3(3): 127?171 ( January 1922). 14 of greatest long-term value to society, emerges out of the passions and interests of the highest-quality scientists asking basic research questions. Rather than defining a particular problem and hiring a scientist who will conduct prescribed research to solve it, I favor identifying the best scientists in very broad areas of endeavor--as defined by their previous research and their publications--and giving them the freedom to define their own research problems and priorities. Restricting individuals of this sort to a narrow specific problem usually leads to narrow findings and less valuable science. Open-ended research by very creative scientists is a better long-term investment strategy. In fact, the Arboretum's capacity to make major advances in our basic understanding of plant biology is unique. There are many institutions with strong agricultural missions (Cornell comes to mind) that are much better positioned to apply massive scientific resources, underwritten by the Department of Agriculture, to the resolution of agricultural or horticultural problems by developing drought- or pest-resistant varieties of crop species, for example. Today, however, the distinction between applied and basic research has become increasingly blurred Senior research scientist David E. Boufford collects herbarium specimens with by the impact of the the help of local Tibetans at an elevation of 3,780 meters, in Daofu Xian county of genomics revolution on western Sichuan, China. In 2007, Arboretum researchers joined an international our understanding of team for the fourth of five expeditions to China's Hengduan Mountain region, home to 30 to 40 percent of China's estimated 30,000 plant species. The group plant physiology and made 2,652 collections of vascular plants for a total of 17,861 herbarium sheets, development. These days and 752 collections of tissue samples for molecular analysis. In addition, members basic research in plant of the team photographed the plants and their habitats for the project's biodiversity website (http:\/\/hengduan.huh.harvard.edu), the largest collection of such biology promises to condocumented and vouchered images of Chinese plants, photo by Susan Kelley. tribute directly to imme- 15 diate societal problems in ways, and in a time frame, that are very hard to predict. Let me cite as examples three major problems that currently confront us and suggest how basic research on woody plants might point to solutions: global climate change, remediation of severely polluted land, and energy independence. Early each year, as the days of spring grow longer, a magical phenomenon sweeps across vast tracts of land in the temperate regions of the world. The branch tips of a wide diversity of deciduous trees develop a covering of carbon-sequestering machines called leaves. Over time, this greening of a large part of the earth's surface has a huge impact on the overall carbon budget of the planet and, therefore, on its climate. Curiously, though, maple trees develop their leaves very early in spring, while for unknown reasons the leaves of oak trees emerge much later. Could basic research on the biology of leaf emergence and canopy development, particularly on the difference in the timing of these phenomena in oaks and maples, contribute to a greater understanding of global climate and how humans are changing it? Consider the second example, remediation of land pollution. There are natural populations of plant species whose roots have adapted to growing in soils that have been severely contaminated by heavy metals, such as mercury. Could an investigation of the basic biology of root growth under these conditions suggest ways that this capacity might be introduced into a fast growing tree such as aspen, thereby allowing the mercury to accumulate in its wood for safe disposal? Finally, the search for renewable energy has increasingly focused on potential fuels from plants (biofuels). Unfortunately, the strong and resilient structure of wood, with which any carpenter is quite familiar, resists the efficient extraction of energy to create a liquid form of fuel able to compete with gasoline. Could a better understanding of the biology of wood lead to a crop that efficiently yields its energy in a highly concentrated form of fuel? In sum, I believe the Arboretum must seize the opportunity before it. We can become an international leader in the type of basic research that will be required to resolve fundamental problems facing today's world. Through the construction of research facilities on Weld Hill, we are taking a giant step toward achieving this goal. M A N A G E M E N T I N I T I AT I V E S F O R COLLECTIONS AND LANDSCAPE T he past five years have brought significant progress in strengthening operations at the Arnold Arboretum. Signaling an increased commitment to excellence in the care of the living collections as well as the landscape in which they grow, staff members completed a strategic plan with the explicit goal of attaining an exemplary level of quality in arboretum management. To define excellence in the care and development of a botanical collection and landscape, Arboretum managers investigated four sister institutions that share our mission as centers of knowledge and investigation relating to woody plants: Holden Arboretum in Kirtland, Ohio; Morris Arboretum of the University of Pennsylvania, Philadelphia; Morton Arboretum, Lisle, Illinois; and the U.S. National Arboretum, Washington, DC. These investigations included interviews with staff, reviews of policies and management practices, and tours of facilities, equipment, and landscapes to explore common challenges and identify best practices across our profession. Arborist Robert Ervin gets a lift to collect seed from the upper branches of an Abies accession for use in the Arboretum's `Tree of Life' investigations of gymnosperms, funded by the National Science Foundation, photo by Kathryn Richardson. In assessing the accomplishments and aspirations of our peer institutions, we identified three key initiatives that will significantly enhance the work of the Arnold Arboretum: Landscape Management, Collections Development, and Plant Health Management. Landscape Management Our careful review of how landscape work is accomplished at sister arboreta underscored the benefits of assigning to each horticulturist the responsibility for specific collections and landscapes. As our professional peers have found, this site-specific focus yields substantial cumulative knowledge, enabling staff to provide increasingly effective horticultural care and to serve as \"local\" experts on soils, pests and disease, collections development, hardscape maintenance, and visitor needs and impacts. In June 2006, following a history of more broadly deploying staff, Arboretum managers implemented this approach through a new Landscape Management Plan. After organizing our 265-acre landscape into 62 management zones composed of contiguous areas that share similar challenges, collections themes, and management priorities, we placed each zone under the care of a staff horticulturist. Now entering its second year of implementation, the new system is yielding substantial improvements, which will be amplified as horticulture manager Steve Schneider collaborates with our dedicated horticulturists to further refine care plans for collections, natural areas, and historic features. In a second phase, to be completed in 2009, the Landscape Management Plan will expand to include curatorial initiatives, cultural resource management goals, and longer-term capital projects. The end result will be a comprehensive vision for the Arnold Arboretum landscape. Collections Development Our assessments also identified a strong need to develop a highly systematic approach to collections development. As we enter our 137th year, the Arboretum collections are distinguished by their maturity, with over 500 accessions that exceed 100 years in age. Through careful planning, the coming decades can bring a significant strengthening of our collections. Priorities for future development include expansion of our national collections in Acer, Carya, Fagus, Syringa, Stewartia, and Tsuga, along with the acquisition of documented, wildcollected accessions to fill critical gaps, or augment specimens of lesser-known provenance. We will also create new research opportunities through increased representation of the disjunct genera of eastern Asia and North America and other taxa that can directly support the work of Arboretum scientists. Additional tasks include assessing space-constrained collections, particularly our plantings of shrubs, as well as implementing new curatorial policies for potentially invasive accessions. In January 2007, Michael Dosmann, a recent graduate of Cornell University's doctoral program in horticultural science, joined the Arboretum staff in the new position of curator of living collections. A former Arnold Arboretum Putnam Fellow, Michael brings strong expertise in both hardy woody plants and collections management. In 2008, he will complete a collections development plan to guide the acquisition of new accessions and to set curatorial priorities for the next five years. Horticultural staff relocates Syringa `Purple Haze' (AA 36-2002) on Bussey Hill from its testing location at the Dana Greenhouse, photo by Steve Schneider. the human-mediated transport of organisms has made the management of hemlock woolly adelgid, winter moth, and other invasive species a challenge of expanding scope. At the same time, we have sought to more effectively manage the complex and changing ecosystem within which our diverse botanical collections grow. To address these and other needs, the strategic vision for horticulture called for a staff position dedicated to overseeing plant health. In the fall of 2007, Julie Coop, former grounds superintendent, was appointed plant health manager. Charged with implementing a comprehensive approach, Julie will coordinate integrated pest management activities and increased monitoring of critical environmental factors over time, including pest populations, soil pH, and soil moisture. This work will provide important opportunities to better understand the impacts of introduced organisms, climatic shifts, and other forces of long-term ecological change in the Arboretum environment. These initiatives promise to bring the Arnold Arboretum to the forefront of professional practice, requiring significant investment but Plant Health Management achieving the standards of excellence and innoNot surprisingly, the larger environmental vation befitting an internationally respected changes altering our world have affected the botanical institution. work of the Arnold Arboretum. Most notably, --Richard Schulhof, deputy director 18 Sargent Fellows E qually important as creating new facilities for research is the development of personnel policies to support the hiring of senior scientists at the Arboretum. Ordinarily at Harvard University faculty members are hired by academic departments to teach while also pursuing scholarly activities. Since the Arboretum is not an academic department--it is administered by the Vice President for Administration rather than being part of an academic school such as the Faculty of Arts and Sciences--we needed to create a new type of position that would attract highly qualified scientists. At the same time, a set of recruitment and retention policies different from those ordinarily associated with our administrative positions would be required. Scientists in a university setting usually have remarkable freedom to define their work schedules and to develop their research programs. The quality of their research is judged according to exactingly high standards set by the larger community of scientists rather than by the administrator who \"supervises\" them. Therefore the Arboretum's personnel policies must be crafted to encourage creativity rather than constrain it. In 2003, working with Harvard botanical faculty members in Cambridge, I defined a new type of position: the Sargent Fellow. Over time, I envision that the Arboretum may have six or eight such Fellows managing their own research programs in the new Weld Hill facility. Initial appointments will be for a fixed term of time (two years, followed after evaluation by another five years). At the end of this time, Fellows will be considered for permanent appointments based on a review of the quality of their research by their peers. In August 2003, the Arboretum appointed the first Sargent Fellow, Dr. Sarah Mathews. Sarah conducts research on the biology and evolution of systems in plants that sense environmental conditions and control plant development. She currently works in borrowed facilities at the Harvard University Herbaria in Cambridge and eagerly anticipates the construction of the Weld Hill facility. In 2004 I recruited a second Sargent Fellow, Dr. Maciej Zwieniecki, who is a plant physiologist interested in the physical and biological mechanisms that control the acquisition and movement of water and nutrients in very large trees. As completion of the Weld Hill facility approaches, I will begin searching for a senior research director, who may also be a faculty member at Harvard. For this position I hope to attract an exceptional scientist who will oversee the development of a broad, long-term research program on the biology of trees. Balancing a Dual Mission I n making this major investment in research, does the Arboretum risk overshadowing its traditional mission to serve the public? For the Arboretum, administered as it is by one of the world's leading research universities, what is the proper balance between the scholarly activities that ordinarily define the mission of Harvard and the programs of education and visitor amenity that serve the general public? During the Arboretum's first half-century, approximately ninety percent of the Arboretum's activity and expenditures would probably have been defined as research by its director, Charles Sprague Sargent. At that time public activities largely consisted of publishing the Bulletin of Popular Information (the predecessor of Arnoldia) and holding occasional public lectures by staff for local schoolteachers. This was also a time when research itself, especially exotic expeditions securing new species in distant lands and introducing them as landscape plants for the garden, was seen by the Arboretum's public--primarily affluent Sargent Fellow Sarah Mathews collects foliage samples of Thuja plicata for DNA analysis as part of the `Tree of Life' project, photo by Mark Beilstein. 20 Bostonians during that period--as more obviously serving their interests (see Current International Research, page 13). Over the last half-century, botanical research, particularly at Harvard, has become much less accessible to non-specialists, while the Arboretum's public has become much more diverse. When I became director in 1989, research as a percentage of the budget was below twenty percent and declining. As described in my most recent Director's Report, in 2002, the scientific reputation of the Arboretum was coasting on the impact of work done decades earlier, and current research activity was being conducted by a few scientists who were housed in Cambridge and distant from the collections here in Boston. If research at the Arboretum was to avoid extinction, a major investment would be required to modernize facilities, expand the scope of research, and fully engage the intellectual resources of the University in a collaborative effort. The Weld Hill facility and our Sargent Fellows program begin to address this need. In the 1980s, at the same time that our research effort was contracting, our public service initiatives, including education programs for children and adults, as well as visitor services, were expanding significantly. Over the past decade and a half, we have increased our commitment to improved care for our collections and landscape (see Management Initiatives for Collections and Landscape, page 16). Large landscape projects, exemplified by the new Leventritt Shrub and Vine Garden, have richly enhanced the enjoyment of visitors. In 2002, we adopted the Landscape Institute (formerly administered by Radcliffe College as the Radcliffe Seminars Landscape Design Program), adding a major public program to our budget. Over the next five to ten years, our investment in research is likely to increase to perhaps sixty percent of our budget, with the remaining forty percent supporting programs that serve the public. This seems about right to me. However, these two parts of our dual mission-- as research institution and as public museum--have always been in some tension at the Arboretum, and they pose an unusual governance challenge for the University. In general, research at Harvard is conducted within its schools, with the directors of research organizations reporting to a dean. This is how the Arboretum was governed from about 1930 to 1988 before Jon Hetman L I N D E N PAT H A N D T H E L E V E N T R I T T G A R D E N I n 2006, completion of a new path through collections of Tilia (linden), Cercidiphyllum (katsura), and Lonicera (honeysuckle) marked an important addition to the Arboretum's pedestrian circulation system. Linden Path, designed by the landscape architectural firm Reed|Hilderbrand Associates, provides a direct connection between Meadow Road and the Leventritt Garden, guiding visitors along a gently winding passage that features exceptional specimens of Tilia and Cercidiphyllum. The Leventritt Garden, also designed by Reed|Hilderbrand, was recognized by the American Society of Landscape Architects with its 2007 Award of Excellence. Completed in 2002 and now a highly popular visitor destination, the Leventritt Garden provides a long-needed home for sun-loving shrubs and vines as well as new interpretive exhibits exploring botanical research, plant conservation, and horticultural introduction. --Richard Schulhof, deputy director S E E D H E R BAR I U M I M AG E PROJ E C T B eginning in the 1960s, Arboretum propagator Al Fordham created a seed herbarium to facilitate the growing of unfamiliar species. Collecting the seed of several hundred rare and unusual taxa, Fordham envisioned a unique resource for the identification and propagation of woody plants from around the world. In 2004, his vision entered the digital age through the Arboretum's Seed Herbarium Image Project (SHIP). Made possible through the generous support of the J. Frank Schmidt Family Charitable Foundation, the Cabot Family Charitable Trust, and the Stanley Smith Horticultural Trust, SHIP uses high-resolution digital photography to document the morphol- ogy of seeds and selected fruit structures. The SHIP images, now available on the Arboretum's website, support scientists, horticulturists, and educators, particularly in propagation research and management of rare and endangered species. The SHIP team is working to finish photographing the Arboretum's six national collections within the North American Plant Collections Consortium: Acer (maple), Carya (hickory), Fagus (beech), Stewartia, Syringa (lilac), and Tsuga (hemlock). Using new protocols and equipment developed for microphotography, SHIP will next document species within the Ericaceae (heath family). --Richard Schulhof, deputy director Seed and fruit images of Stewartia ovata var. grandiflora created as part of the Seed Herbarium Imaging Project (SHIP) include a detail of a single seed, multiple seeds displaying alternate views and morphological variation, seeds shown with fruiting structure, and fruiting detail, photos by Julie McIntosh Shapiro. 23 it was moved to administration within the Central Administration. At about the same time, other non-research organizations providing public service, like the University Art Museums and the American Repertory Theatre, were also moved to the Central Administration. Today, with the revitalization of our research mission and our major investment in research facilities and staff, we have become something of an anomaly in Central Administration and this creates some specific problems for the Arboretum. First, there are no personnel policies for research scientists in the Central Administration of the University because research scientists at Harvard ordinarily receive academic appointment in one of the schools. Second, the Arboretum now manages seven federal research grants-- another anomalous activity within Central Administration--and the number promises to grow. Finally, because neither the director of the Arboretum nor the researchers on its staff are members of a faculty, there is no link between long-range planning for research at Harvard, which happens in faculty meetings and committees, and long-range planning at the Arboretum which does not involve faculty. The issue of governance for the Arboretum is further complicated by the administrative position of the Harvard University Herbaria (HUH), located five miles away in Cambridge. Nearly half the herbarium and library collections within this botanical unit are the property of the Arnold Arboretum and we pay for about 40% of the facility's operation. Yet its director reports through a school to the Dean of the Faculty of Arts and Sciences (FAS) and the administrative and personnel policies governing its operation are defined and executed by FAS.3 Despite considerable similarity in their missions, these two botanical units are governed in very different parts of the University. All of these factors have come together to frame a critical question: Should the Arboretum return to management by the Faculty of Arts and Sciences, consistent with its research mission? Or should it remain under the management of Central Administration, consistent with its public mission? This matter should be resolved before a new director is chosen for either the Arboretum or HUH, but it presents a thorny challenge. In the case of the Arboretum, a governance structure that is dominated 3 In July, 2005, I accepted a three year term as director of HUH in addition to my duties as director of the Arboretum. LANDSCAPE INSTITUTE S ince its relocation to the Arnold Arboretum in 2002, the Landscape Institute (formerly the Radcliffe Seminars Landscape Design Program) has undergone several changes that will prove critical to its future success as an educational resource for professionals working in landscape design and management, historic preservation, landscape history, and related fields. In 2006, the Institute moved from the Cronkhite Center, the program's home for more than seven years, to 29 Garden Street in Cambridge, just outside Harvard Square. The new facility, with its strikingly contemporary design, offers increased space and new possibilities for classes and special programs. An equally significant transition began in early 2007 when Landscape Institute director John Furlong, following twenty-five years of strong leadership, announced his desire to step down to devote more time to private practice and teaching. Following a national search, Heather Heimarck assumed full-time duties as director in February 2008. Her background includes a Master in Landscape Architecture from the Harvard Graduate School of Design, work with several accomplished landscape architects, and the founding of Heimarck and Foglia, formerly HighMark Land Design, a firm specializing in green design, innovative use of plant materials, and new construction approaches. In keeping with the progressive spirit of the Institute, Heather's extensive practical experience and commitment to innovation promise important new directions in sustainable design, construction, and landscape horticulture. Looking to the future, the Landscape Institute will continue the work of Arboretum founding director Charles Sprague Sargent to strengthen the landscape professions, while also addressing burgeoning societal needs for leadership in environmental design and management. --Richard Schulhof, deputy director by either FAS or Central Administration risks undermining one part of our dual mission or the other. Put another way, if the dual mission of the Arboretum is to survive into the future, its governance structure must support integration of its research investment with the traditional mission of the University (research and undergraduate education) while also encouraging the public mission of the Arboretum to thrive. A New Model for the Arboretum T hus our sizable investment in research will only succeed for the Arnold Arboretum if it also succeeds for the University. Effectively this means that the conduct of research must be governed as an integrated part of ordinary academic operations and coordinated through members of a school's faculty. In this case, the logical school is FAS, where the Arboretum resided prior to 1988. I believe the decision about how Arboretum research is to be managed will need to recognize this reality. The remaining question about Arboretum governance regards its public mission: what threats might it face in an academic context such as FAS? Magnolia `Elizabeth' (AA 120-78), photo by Jim Harrison 26 Three come to mind. First, a large bureaucracy exists within FAS to manage intelligent and ambitious professors who are powerful through their appointment and their relations with outside sources of funds (donors, government agencies, consultancies). Professors, in turn, are always creative in their efforts to circumvent a constraining bureaucracy. A small public museum governed by this sort of bureaucracy without the protection of a resident professor can suffocate through no one's ill intentions. Second, public service will never be a high priority at major research universities beyond the normal public relations activity required by proximity to non-university neighbors. Put more positively, universities like Harvard provide \"public service\" for the long term: they develop tomorrow's leaders and increase our understanding of the world. In such a university, a unit providing direct services to the public can suffer from administrative and financial neglect simply because it is not a priority for senior administrators and deans. Finally, unprotected public service operations in the midst of large academic schools can suffer financial predation from wellmeaning but narrowly focused academics. Funds that are not being used to educate university students or support faculty research look to many professors like money being poured into a hole, money that could be more fruitfully spent on academic activities. In view of these threats, three elements seem essential to protecting a university unit with a mission of direct public service--what we here call the public museum of the Arboretum. First, the public museum must have a clear identity and sufficient independence to establish its own brand of administrative culture, one that openly acknowledges a commitment to service. This requires real authority, particularly over budgets and personnel, even though in an academic setting this authority is normally vested in the dean and his or her administration. Second, this independence can only work if the public museum is given a guaranteed base of financial support sufficient to carry on essential activities. This base of support must be ensured a reasonable rate of annual increase to sustain it against inflation in the future. While this will support the core program, special projects that enhance its facilities and programs will also require the public museum to raise money from the public, largely through philan- ENHANCING VISITOR EXPERIENCE A Time for Change, a strategic vision authored by Bob Cook in 2002, voiced a strong commitment to improving the quality of information and orientation furnished to Arnold Arboretum visitors. After several years of planning, 2007 brought completion of a comprehensive wayfinding signage system that enables visitors to fully explore our 265-acre historic landscape and diverse botanical collection. Created by the environmental graphic design firm Roll Barresi, the system provides \"you are here\" maps, path markers, and other navigational aides to ensure confident and effective visitor wayfinding. In 2009, the system will be augmented by a new map brochure and temporary interpretive signs focused on seasonal information. To provide leadership for visitor programs, Julie Warsowe, a graduate of Cornell University's program in public garden management, was appointed the Arboretum's first manager of visitor education in 2006. Julie recently completed a survey of Arboretum visitors that provides valuable information about the demographics, motivations, and interests of the broad community that utilizes our landscape. Survey data, compiled and analyzed by visitor research consultants People, Places and Design, will inform development of new interpretive programs, including plans for new exhibits in the Hunnewell Building Visitor Center. Justin Ide\/Harvard News Office, ?2004 President and Fellows of Harvard College D I G I TA L R E S O U R C E S Plant Collections ? Plant Inventory: Search the Arboretum's living collections database by common or scientific name; http:\/\/arboretum.harvard.edu\/plants\/inventory.html ? Interactive Map of the Arboretum: Explore 31 plant collections and 76 featured plants; http:\/\/www.arboretum.harvard.edu\/visitors\/map.html?myURL=\/visitors\/visitors.html&myLayer=collections ? Multisite Plant Inventories: Search 24 living collections and conservation databases for participating botanical institutions, hosted by the Royal Botanic Garden, Edinburgh, Scotland, UK; http:\/\/rbg-web2.rbge.org.uk\/multisite\/multisite3.php Herbarium Collections ? Cultivated Herbarium: Search the Arboretum's herbarium collections by common or scientific name; http:\/\/arboretum.harvard.edu\/plants\/herbarium.html ? Seed Herbarium: Browse images from the Arboretum's seed herbarium; http:\/\/arboretum.harvard.edu\/plants\/herbarium.html ? Joseph Rock's Type Specimens: Access to 197 type specimens collected by Joseph Rock between 1923?1932 in western China and Tibet; http:\/\/www.arboretum.harvard.edu\/library\/tibet\/herbarium.html ? Maps of Joseph Rock: Navigate 10 individual hand-drawn maps and related gazetteer illustrating plant explorer Joseph Rock's travels (1924?1927) in China; Maps: www.arboretum.harvard.edu\/library\/tibet\/zoom\/rock_maps.html Gazetteer: www.arboretum.harvard.edu\/library\/tibet\/map.html Photographic Collections ? The Arboretum Through Time: Historical photographs of the Arboretum's landscape and collections; www.arboretum.harvard.edu\/programs\/views\/intro.html ? Botanical and Cultural Images of Eastern Asia, 1907?1927: Archival images of Arboretum plant explorers and their field photographs from their exhibitions; www.arboretum.harvard.edu\/programs\/eastern_asia\/overview.html ? South Central China and Tibet: Hotspot of Diversity: Images of natural history and ethnographic collections from Arnold Arboretum expeditions to China and Tibet between 1924 and the present; http:\/\/www.arboretum.harvard.edu\/library\/tibet\/expeditions.html ? Cienfuegos Botanical Garden, Cuba: Archival and contemporary photographs of the former Atkins Institution, administered by the Arnold Arboretum from 1946 to 1959; http:\/\/www.arboretum.harvard.edu\/programs\/cuba\/intro.html Other Resources ? Arnoldia: Searchable database of all volumes published during the last hundred years of the Arboretum's journal Arnoldia and its antecedent, The Bulletin of Popular Information; http:\/\/arnoldia.arboretum.harvard.edu ? Silva: All issues of the Arboretum's news magazine since its inception in 2005; http:\/\/arboretum.harvard.edu\/aboutus\/silva\/current.html ? Correspondence Index: A work in progress referencing institutional correspondence from the 1880s to 1940; http:\/\/www.arboretum.harvard.edu\/library\/about_arc.html ? OASIS: Harvard's Online Archival Search Information System includes 42 finding aids to archival and manuscript records at the Arnold Arboretum Archives; http:\/\/oasis.harvard.edu ? Hollis: Searchable database containing more than 9 million records for more than 15 million items in the Harvard University Libraries; http:\/\/lib.harvard.edu\/ ? Google Book Search: Search the full text of all books available in Google Book Search, including some 3,000 titles from the Arnold Arboretum Library; http:\/\/books.google.com 29 thropy rather than earned revenue which usually creates too many conflicts in an academic culture. Finally, a public museum in academia needs deeply committed leadership with a strong belief in its public mission, not only within its internal operation but also at more senior levels of the university. For the Arboretum, this certainly includes the director, who may be primarily focused on the health of the research mission. But it must also extend to the supervising administrator, whether in FAS (the dean) or in Central Administration (the provost or a vice president). These governance issues are now being addressed in two ways. First, the provost has formally assigned them for deliberation to the University's senior committee for long-range planning for science and engineering. I anticipate resolution of the matter by the end of the fiscal year in June. Second, at the Arboretum we have begun to develop a new model of operation that more explicitly acknowledges the identity of our public museum functions traditionally serving our visitors and students. The Public Museum D efining the programs at the Arnold Arboretum that provide public service is relatively easy in the case of educational programs and visitor services. Our educational programs are of two kinds. Children's education serves about 2,000 individuals under the age of ten each year. Adult programs include lectures and classes for the dedicated amateur and the more formal programs of the Landscape Institute, which can lead to a certificate indicating professional-level achievement. The services for visitors might be characterized as providing access to information. They include the traditional functions of our horticultural library and our visitor's center, both of which serve people walking in through the door. But they have also been expanded to include the vast amount of information we are now making available through the Internet (see Digital Resources, page 28). Because digital access provides valuable information to anyone anywhere in the world at any time, these efforts promise to continue growing in the years to come. SCHOOL PROGRAMS ver the course of several years, Arboretum friends Henry and Nod Meyer have generously supported explorations of the natural environment for urban children. Thanks to their encouragement, Arboretum field study programs have grown significantly in depth and focus, now hosting over 2,000 A second initiative serves pre-schoolers attending neighborhood Head Start centers. Over the past three years, visiting children from Hyde Park, Jamaica Plain, and Roslindale were led by trained volunteers in exploring flowers, fruit, bark, soil, and other aspects of plants and landscape. Designed in collaboration with specialists in early children annually with a special commitment to elementary schools in Boston and Chelsea, and to local Head Start centers. Among these initiatives is a new field study that invites Boston students to investigate the dramatic changes caused by hemlock woolly adelgid on the Arboretum's Hemlock Hill. The new program, developed in collaboration with Boston teachers and titled \"A Changing Ecosystem,\" supports the fifth-grade science curriculum through a case study in biological invasion that powerfully conveys the dynamics of ecological change. childhood education, the program provides children with a range of new outdoor experiences, builds vocabulary, and encourages curiosity about the natural world. In coming years, we will increase our training for Head Start instructors in inquiry-based teaching methods and also offer family activities that foster parental participation in pre-school education. In addition to supporting public education, these efforts seek to strengthen neighborhood connections and promote greater enjoyment of the Arboretum by the diverse communities of Boston. --Richard Schulhof, deputy director Kris Snibbe\/Harvard News Office, ?2007 President and Fellows of Harvard College O 31 One might ask, does the Arboretum's program in horticulture also provide \"public service\"? While the living collections are of great value for research, they and the grounds are given a much higher level of care than would be the case if they were grown solely for the use of scientists. A planting for purely scientific use would look like an agricultural field, with long rows of trees enclosed by a tall fence, rather than an open, beautiful landscape inviting the visitor to linger. I therefore see our strong commitment to horticultural excellence on the grounds as a public service, though that commitment is shared with our obligation to serve the needs of researchers today and tomorrow. This dual function of the living collections is the essence of Charles Sprague Sargent's original vision for the Arboretum. Last summer we reorganized the Arboretum into two programmatic entities, research and public museum, supported by a central administration consisting of the director's office and the functions of finance, information technology, and facilities (see Organizational Chart, page 32). As noted earlier, the research program will be headed by a director of research and will include all senior research staff, as well as the post-doctoral fellows and technicians in their laboratories. The public museum will include the functions of horticulture, education, and information access for visitors. It is now headed by my deputy director, Richard Schulhof. Anticipating its need for financial security, we have identified a core budget for the public museum that for the current fiscal year amounts to $9,375,000 (out of a total Arboretum budget of $13,770,000; see Summary of Operations, page 33). We have also projected a four-percent rate of budget increase for future years. Finally we have transferred the Arboretum's membership and development program from administration (reporting to the director) to the public museum (reporting to the deputy director). Horticultural technologist Scott Grimshaw cuts invasive undergrowth of Rubus spp. growing along Oak Path, photo by Richard Schulhof. 32 The Arboretum and its Friends ne may question the change in the reporting relationship of the development department. Shouldn't it support the entire institution, not solely the public museum? Ever since its creation in 1872, the Arboretum has benefited from very generous friends and their generosity has arrived with remarkably few restrictions beyond a specification that their gifts should support the Arboretum (as opposed to Harvard University). I believe this pattern of giving reflects unusual trust on the part of our friends in the wisdom of past directors. O AR N O LD AR B O R E TU M O F H ARVAR D U N I V E R S IT Y O R G A N I Z AT I O N A L C H A R T President of Harvard University Drew Gilpin-Faust Dean of the Faculty of Arts and Sciences Vice President for Administration Michael D. Smith Sally Zeckhauser Director of the Harvard University Herbaria Director of the Arnold Arboretum of Harvard University Bob Cook Bob Cook Director of Research Director of Capital Projects (TBH) Cynthia Jensen Sargent Fellows Education & Public Programs Horticulture Deputy Director Richard Schulhof Development Finance Director of Human Resources Director of Administration & Finance Lisa Toste Andrea Nix IT Library Facilities S U M M A R Y O F O P E R AT I O N S T his chart indicates our overall financial performance for the past six years. As we have taken on new capital projects and invested in research, our overall budget has moved from overall surpluses to deficits in the past two years which have been covered by reserves. With respect to income trends, the appointment of Sargent Fellows after 2003 has led to increases in research grants, and renewed efforts in membership have brought in increasing numbers of gifts. The large jump in education income in 2003 represents the assumption of administrative responsibility for the Landscape Institute and its educational programming; it is offset by equal increases in expenditures. With respect to expenses, salaries and services are the major categories increasing, along with debt service reflecting the completion of the Leventritt Garden in 2005. Expenses will continue to increase in coming years as we expand our investment in research. Actual FY 2002 Actual FY 2003 Actual FY 2004 Actual FY 2005 Actual FY 2006 Actual FY 2007 7,666,661 8,226,848 8,244,215 8,360,913 9,042,906 9,590,265 Membership\/Gifts 644,972 497,892 376,947 292,612 307,663 442,613 Enterprise 116,774 115,463 323,496 239,770 334,377 346,373 Grants 112,446 39,422 194,647 410,335 455,040 566,553 70,799 605,748 607,701 675,098 753,989 636,561 8,611,652 9,485,373 9,620,006 9,845,844 10,754,143 11,434,580 4,162,438 4,393,437 5,000,632 5,801,809 6,219,559 6,526,842 Supplies\/Equipment 429,101 460,973 554,039 560,997 600,118 562,473 Facilities\/Operations 717,645 881,689 724,302 800,482 1,143,790 1,505,463 Services 771,907 1,058,734 1,296,032 1,131,235 1,931,237 1,990,992 University Subvention 256,483 264,092 292,123 317,134 346,520 378,988 66,934 107,962 142,577 235,863 182,432 201,637 6,404,508 7,166,887 7,882,705 8,714,636 10,283,824 11,018,610 2,207,144 2,318,486 1,737,301 1,131,208 470,319 415,970 312,416 304,434 393,609 609,450 566,729 568,962 1,894,728 2,014,052 1,343,692 521,758 (96,410) (152,992) Income Endowments Education\/Publications Total Income Expenses Salaries\/Benefits Travel Total Expenses Excess (Loss) Debt Payment Total Excess (Loss) 34 The research program of the Arboretum, now and as expanded in the near future, will be fully integrated into the University's overall research endeavors in the botanical sciences. As such it will benefit from the fundraising energies of the University, which are traditionally focused on alumni. Fundraising for the public museum will face the significant challenge of operating nimbly in the shadow of Harvard's larger fundraising program which is unlikely to support public service as a priority. Because most members of the Arboretum are unusually committed to the horticultural and educational work of the institution, they give their support to ensure the continuing excellence of these programs. Therefore I believe that the work of our development staff should primarily sustain the long-term survival and growth of the public mission. So a bargain can be struck. The University will commit itself to ensuring the health and well-being of the Arboretum's research mission, at the same time permitting the Arboretum to raise funds for its parallel mission of public service. But the Arboretum will need help achieving this second goal. Our members and friends, who have always been deeply loyal to our public purposes, will be asked to increase their critical support for this equally important part of our mission. It is a bargain that can secure the continuing fulfillment of Charles Sargent's remarkable vision. Beech Path, photo by Eric Roth Robert E. Cook 1 May 2008 A R N O L D A R B O R E T U M W E AT H E R S TAT I O N DATA --20 07 Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct. Nov. Dec. Avg. Max. Temp. (?F) Avg. Min. Temp. (?F) Avg. Temp. (?F) Max. Temp. (?F) Min. Temp. (?F) Precipitation (in.) Snowfall (in.) 40.0 33.0 46.4 52.8 71.4 77.0 81.7 81.7 76.6 66.8 50.3 36.4 23.1 16.1 25.8 36.4 49.5 57.4 63.6 61.2 55.9 47.9 31.5 20.5 31.55 24.55 36.1 44.6 60.45 67.2 72.65 71.45 66.25 57.35 40.9 28.45 69 49 70 87 93 95 91 95 94 87 68 54 2 7 4 27 33 47 53 50 43 32 20 8 2.74 2.15 4.78 7.91 3.16 3.1 3.44 0.4 1.67 2.59 3.11 5.93 1.5 7 7.7 0 0 0 0 0 0 0 0 26.7 Avg. Maximum Temperature Avg. Minimum Temperature Avg. Temperature Total Precipitation Total Snowfall Warmest Temperature Coldest Temperature Last Frost Date First Frost Date Growing Season 59.5? 40.7? 50.13? 40.98\" 42.9\" 95? on August 4 2? on January 26 and 27 32? on April 9 32? on October 9 183 days 2007 was a year of highly changeable weather at the Arnold Arboretum. The first half of January brought record warmth, even inducing a scattering of cherries into early bloom. But on January 16 winter finally arrived in the Northeast and temperatures plummeted. The first 16 days of January averaged 12 degrees above normal while the second half of the month averaged 5 degrees below normal. The winter cold extended well into March and April and was coupled with heavy rains and street flooding during both months. Spring and early summer precipitation provided adequate levels of moisture for a very good growing season up to the end of July, which began a period of extremely dry weather in the Northeast. Fortunately temperatures did not soar during this dry spell which helped to limit the negative impact of the drought. However it was dry enough to delay and significantly reduce fall planting from Arboretum nurseries to the grounds. September and October were pleasantly mild and quite sunny. Although poor fall color had been predicted (in part because of the drought), the foliage at the Arboretum ended up putting on quite a colorful show. November remained on the dry side and brought the onset of winter with below average temperatures. To close the year, December was very cold and snowy, with snow accumulation coming close to breaking the record for Boston. Unlike the very mild late fall\/early winter of 2006, this period in 2007 reminded us how tough our winters can be. Throughout this variable year of weather, visitors and staff continued to appreciate the beauty of the Arnold Arboretum through all the seasons. 36 S TA F F O F T H E A R N O L D A R B O R E T U M * ADMINISTRATION Rose Balan, Staff Assistant Donna Barrett, Accounting Assistant Kenneth Clarke, Horticultural Technologist (resigned 9.30.05) Robert Cook, Director, Arnold Professor Ann Marie Countie, Computer Services Manager William Hays, Biological Database Applications Programmer (resigned 6.3.06) Andrew Hubble, Network Systems Manager Cynthia Jensen, Director of Capital Projects (hired 1.3.05) James Macklin, Director of Collections and Informatics (hired 3.20.06) Frances Maguire, Director of Administration and Finance Andrea Nix, Director of Administration and Finance (hired 4.30.07) Karen Pinto, Staff Assistant Christopher Preheim, Executive Assistant (hired 9.11.05) David Russo, Facilities Manager Lisa Toste, Director of Human Resources (hired 7.1.02) Sylvia Winter, Landscape Project Manager (hired 9.2.03) DEVELOPMENT\/INSTITUTIONAL ADVANCEMENT Sheila Baskin, Development Assistant Anne Jackson Bell, Events Manager (resigned 7.27.06) Ronda Brands, Development Assistant (hired 7.1.05 and resigned 1.5.07) Jon Hetman, Development Manager Wendy Krauss, Development Assistant (hired 1.24.07) Michele Levy, Director of Communications (hired 2.6.07) Julie Anne McNary, Senior Development Officer (hired 2.6.05 and resigned 8.26.05) Heidi Norris, Development Officer (hired 2.7.06) Robert Surabian, Director of Development (hired 8.9.04) Michaela Tally, Events Manager (hired 2.26.07) *(1 July 2002 through 30 June 2007) PUBLIC & PROFESSIONAL PROGRAMS Sheryl Barnes, Webmaster Kirstin Behn, Staff Assistant (resigned 6.17.05) Ellen Bennett, Manager of Horticultural Information (resigned 1.10.03) Sonia Brenner, Staff Assistant (resigned 6.1.07) Lois Brown, Editorial Assistant (hired 9.25.05) John Furlong, Director of the Landscape Institute (program transferred from Radcliffe 7.1.02) Ann-Marie Greaney-Williams, Administrative Coordinator (hired 7.1.02) Leah Kane, Staff Assistant (hired 5.2.05) Karen Madsen, Editor of Arnoldia Sandra Morgan, Staff Assistant (resigned 5.3.07) Caroline Richardson, Manager of Horticultural Information (hired 5.19.03 and resigned 12.15.05) Nancy Sableski, Manager of Children's Education Micah Schatz, Arboretum Assistant (less than half time); (hired 3.27.04 and resigned 4.22.07) Richard Schulhof, Deputy Director (hired 9.30.02) Pamela Thompson, Manager of Adult Education Julie Warsowe, Manager of Visitor Education (hired 7.5.06) Sheryl White, Staff Assistant Laura Wilson, Staff Assistant (hired 7.10.02) HERBARIUM David Boufford, Senior Research Scientist Maria del Carmen Chavez-Ortiz, Curatorial Assistant (hired 8.1.05 and resigned 7.31.06) Lihong (Wendy) Duan, Curatorial Assistant (transferred to J.P. Library 1.2.05); Staff Assistant (hired 10.31.05) Jennifer Fonda, Curatorial Assistant (hired 11.9.04) Susan Hardy Brown, Curatorial Assistant Edith Hollender, Arboretum Assistant (less than half time) Henry Kesner, Curatorial Assistant (hired 10.5.04) Walter Kittredge, Curatorial Assistant Jude Mulle, Curatorial Assistant (resigned 9.30.02) Melanie Schori, Editorial Assistant (resigned 10.10.03) Emily Wood, Manager of Systematic Collections 37 LIBRARY Beth Bayley, Library Assistant (hired 5.1.04) Sheila Connor, Horticultural Research Archivist Carol David, Library Assistant (resigned 6.1.04) Lihong (Wendy) Duan, Staff Assistant (transferred from Herbarium 1.2.05) Marla Gearhart, Library Assistant (hired 11.29.04) Judy Green, Project Image Cataloger (hired 4.1.01 and resigned 6.30.02) Joseph Melanson, Library Assistant (resigned 12.6.04) Lisa Pearson, Library Assistant (hired 7.8.02) Cathleen Pfister, Library Assistant Christy S. Robson, Catalog Librarian Gretchen Wade, Reference\/Collection Development Librarian Judith Warnement, Librarian of Harvard University Botany Libraries Winifred Wilkens, Library Assistant (retired 9.6.03) LIVING COLLECTIONS Thomas Akin, Assistant Superintendent of Grounds (resigned 10.24.03) John Alexander, Plant Propagator James Allen, Arboretum Assistant (less than half time) Jesse Batty, Grounds Crew Term (9.4.05?9.21.06) Stacy Berghammer, Apprentice (resigned 12.2.02) Jessica Blohm, Gardener (hired 9.10.04 and resigned 7.15.06) Laura Tenny Brogna, Landscape Project Manager (resigned 7.19.04) Julie Coop, Manager of Horticulture John DelRosso, Head Arborist Peter Del Tredici, Director of Living Collections (transferred to Research 7.1.03) Kristin DeSouza, Apprentice (hired 8.29.04 and resigned 9.19.05) Michael Dosmann, Curator of Living Collections (hired 1.2.07) James Doyle, Gardener\/Arborist Apprentice (hired 9.29.03 and resigned 1.5.07) Ralph Ebener, Grounds Crew Term (10.3.04?10.21.05) Charlotte Enfield, Grounds Crew Term (10.30.05? 4.29.06) Robert Ervin, Arborist (hired 7.8.02) David Falk, Horticulture Term (9.5.06?1.5.07) Robert Famiglietti, Horticultural Technologist Kirsten Ganshaw, Horticultural Technologist Donald Garrick, Horticultural Technologist (resigned 7.8.03) Bethany Grasso, Horticultural Technologist (resigned 1.14.04) Scott Grimshaw, Horticultural Technologist (hired 5.17.04) Dennis Harris, Horticultural Technologist Eric Hsu, Putnam Fellow (hired 10.31.05 and resigned 9.23.06) Irina Kadis, Curatorial Assistant Wesley Kalloch, Horticultural Technologist (hired 4.24.06) Susan Kelley, Curatorial Assistant (transferred to Research 1.1.04) Jennifer Kettell, Horticultural Technologist (hired 11.17.03) Alice Kitajima, Apprentice (hired 9.30.02 and resigned 9.21.03) Jianhua Li, Botanical\/Horticultural Taxonomist (transferred to Research 1.1.04) Daniel March, Apprentice (resigned 8.23.02) Brendan McCarthy, Horticultural Technologist (hired 3.20.07) Bruce Munch, Horticultural Technologist Chloe Nathan, Grounds Crew Term (9.4.05?3.3.06) James Nickerson, Horticultural Technologist (resigned 10.8.04) James Papargiris, Horticultural Technologist, appointment as Working Foreperson Thomas Por, Arborist (resigned 9.9.05) Kyle Port, Manager of Plant Records Chris Rice, Horticultural Technologist (hired 6.1.04 and resigned 10.28.05) Kathryn Richardson, Curatorial Assistant (hired 6.7.04) Kelly Ruth, Horticulture Term (9.3.06?1.26.07) Nima Samimi, Gardener (hired 2.16.07) Stephen Schneider, Associate Manager of Horticulture Rita Schwantes, Grounds Crew Term (10.3.04? 11.15.04) Julie Shapiro, Curatorial Assistant (hired 4.9.06) Maurice Sheehan, Horticultural Technologist, Working Foreman (retired 10.31.03) 38 Kyle Stephens, Arborist Apprentice\/Arborist (hired 10.2.06) Kevin Stevens, Apprentice (hired 9.13.05 and resigned 7.28.06) Sara Straate, Curatorial Assistant (resigned 9.16.02) Siobhan Sullivan, Horticulture Term (9.3.06?3.3.07) Aneiage Van Batenburg, Apprentice (hired 9.29.03 and resigned 8.31.04) Mark Walkama, Horticultural Technologist Thomas Ward, Manager of the Greenhouse Victoria Woodruff, Gardener (hired 9.29.03 and resigned 8.5.04) INSTITUTE FOR CULTURAL LANDSCAPE STUDIES (incorporated into Public and Professional Programs) Phyllis Andersen, Director of the ICLS (retired 6.30.04) RESEARCH Kobinah Abdul-Salim, Mercer Fellow (appointed 12.1.02?5.31.03) Jennifer L. Baltzer, CTFS-AA Asia Post Doctoral Fellow (appointed 4.4.05?3.31.2007) Mark Beilstein, Mercer Fellow (hired 12.18.07) Jonathan Bennett, Research Fellow (appointed 8.1.03? 7.31.04) Tim Brodribb, Putnam Fellow (appointed 6.1.05) Zhiduan Chen, Mercer Fellow (appointment ended 8.31.02) Stuart Davies, Science Director of the CTFS-AA Asia Program (resigned 9.30.05); Director of Asian Programs (re-hired 7.1.07) Peter Del Tredici, Senior Research Scientist (transferred from Living Collections 7.1.03) Michael Dosmann, Putnam Fellow (appointment ended 8.31.02) Rodger Evans, Mercer Fellow (hired 1.2.07 and resigned 4.30.07) Kenneth Feeley, CTFS-AA Asia Post Doctoral Fellow (appointed 6.1.05) Margaret Frank, Research Assistant (hired 6.18.07) Lianming Gao, Mercer Fellow (hired 6.26.07) Phyllis Glass, Staff Assistant (hired 5.27.03 and resigned 6.1.05) Anna Gorska, Post Doctoral Fellow (hired 3.1.06) Barbara Gravendeel, Mercer Fellow (hired 10.19.05 and resigned 10.18.06) Jocelyn Hall, Mercer Fellow (appointed 9.1.03?1.31.06) Maria Jaramillo, Mercer Fellow (appointment ended 6.30.03) Zhen Jiao, Mercer Fellow (hired 3.20.06) (resigned 9.14.06) Susan Kelley, Botanical Project Manager (transferred from Living Collections 1.1.04) David King, CTFS-AA Asia Post Doctoral Fellow (appointed 1.1.03?12.31.04) Jeremy Ledger, Research Assistant (hired 7.15.02 and resigned 1.23.04) Ethan Levesque, Research Assistant (hired 12.2.03 and resigned 5.23.07) Jianhua Li, Senior Research Scientist (transferred from Living Collections 1.1.04) Wenbo Liao, Mercer Fellow (appointed 3.1.05?8.15.05) Stuart Lindsay, Mercer Fellow (appointment ended 9.30.02) Tatyana Livshultz, Mercer Fellow (appointed 7.16.03? 10.15.05) Laura Lukas, Arboretum Assistant (hired 1.7.05, less than half time); Research Assistant (hired 10.9.05 and resigned 1.14.06) Andrew Marshall, Mercer Fellow (appointed 8.1.04? 8.1.06) Sarah Mathews, Sargent Fellow (appointed 8.11.03) Joel McNeal, Post Doctoral Fellow (appointed 2.22.05? 1.26.07) David Middleton, Tropical Plant Systematist (resigned 12.31.04) Rebecca Pradhan, Mercer Fellow (appointed 9.1.02? 6.30.03) Richard Primack, Putnam Fellow (hired 7.1.06) Hardeep Rai, Post Doctoral Fellow (hired 5.1.07) Sabrina Russo, CTFS-AA Asia Post Doctoral Fellow (appointed 9.1.03?12.31.05) Lawrence Sack, Putnam Fellow (appointed 8.1.02? 7.31.03) Sonali Saha, Putnam Fellow (appointed 10.8.02? 8.31.04) Mariya Schilz, Research Assistant (hired 8.20.06) Suzie Shoup, Research Assistant (hired 3.29.04 and resigned 11.13.06) Stephanie Stuart, Research Assistant (hired 11.28.04 and resigned 8.4.05) 39 Wayne Takeuchi, Tropical Forest Biologist Nina Theis, Putnam Fellow (appointed 1.12.04? 8.31.05) Donna Tremonte, Research Assistant (hired 1.5.04 and resigned 8.31.06) Sonia Uyterhoeven, Putnam Fellow (appointed 1.1.02? 2.1.03) Ellen VanScoyoc, Staff Assistant (resigned 6.30.03) James E. Watkins, Mercer Fellow (hired 9.1.06) Campbell Webb, Tropical Forest Biologist (hired 9.1.05) Kyle Williams, Post Doctoral Fellow (appointed 1.12.04?1.12.07) Qing Ye, Post Doctoral Fellow (hired 3.27.06) Jipei Yue, Mercer Fellow (appointed 10.1.04?5.31.05) Lihua Zhou, Putnam Fellow (appointed 10.1.02? 3.31.03) Maciej Zwieniecki, Sargent Fellow (appointed 6.6.04) RESEARCH AFFILIATES Glenn Steven Adelson, Arnold Arboretum Associate (appointed 9.1.06) Ihsan Al-Shehbaz, Arnold Arboretum Associate (appointed 4.1.05) Phyllis Andersen, Arnold Arboretum Associate (appointed 8.1.04) Peter Ashton, Charles Bullard Professor of Forestry, emeritus Jennifer Baltzer, Arnold Arboretum Associate (appointed 4.1.07) Mark Beilstein, Arnold Arboretum Associate (appointed 8.8.06?12.17.07) Gordon Burleigh, Arnold Arboretum Associate (appointed 7.19.06?10.31.06) Mabel Cabot, Arnold Arboretum Associate (appointed 11.1.05) Thomas Campanella, Arnold Arboretum Associate (appointment ended 1.31.03) Chin-Sung Chang, Arnold Arboretum Associate (appointed 9.15.06) Chua Siew Chin, Arnold Arboretum Associate (appointed 3.1.06) Stuart Davies, Arnold Arboretum Associate (appointed 10.1.05; hired 7.1.07) Michael Dosmann, Arnold Arboretum Associate (appointed 4.1.04?3.31.06; hired 1.2.07) Peter J. Franks, Arnold Arboretum Associate (appointed 12.1.02?11.30.03) Irwin L. Goldman, Arnold Arboretum Associate (appointment ended 1.31.03) Jocelyn Hall, Arnold Arboretum Associate (appointed 2.1.06?6.30.06) Richard Howard, Professor of Dendrology, emeritus (died 9.18.03) Shiu-Ying Hu Hsu, Botanist, emerita Alice Ingerson, Arnold Arboretum Associate (appointed 7.1.02?6.30.05) Yu Jia, Arnold Arboretum Associate (appointed 1.1.06) David King, Arnold Arboretum Associate (appointed 1.1.05?9.30.05) James LaFrankie, Arnold Arboretum Associate (appointed 9.1.02?8.31.05) Timothy Laman, Arnold Arboretum Associate (appointment ended 6.30.04) Tatyana Livshultz, Arnold Arboretum Associate (10.16.05?12.31.05; hired 3.1.06) Richard H. Ree, Arnold Arboretum Associate (appointed 2.1.03) Kenichi Shono, Arnold Arboretum Associate (appointed 1.1.05 ?12.31.05) Stephen Spongberg, Curator, emeritus George Staples, Arnold Arboretum Associate (appointed 8.1.03?7.31.04) Hang Sun, Arnold Arboretum Associate (appointed 1.1.06) Kim Tripp, Arnold Arboretum Associate (appointment ended 4.30.03) Sonia Uyterhoeven, Arnold Arboretum Associate (appointed 2.1.03?1.31.05) Keith Vanderhye, Arnold Arboretum Associate (appointed 9.1.03?8.31.04) Campbell Webb, Arnold Arboretum Associate (appointment ended 8.31.05) Kyle Williams, Arnold Arboretum Associate (appointed 1.13.07) Carroll Wood, Jr., Professor of Biology, emeritus Zhuliang Yang, Arnold Arboretum Associate (appointed 1.1.06) Donglin Zhang, Arnold Arboretum Associate (appointment ended 8.31.02) 40 PUBLISHED WRITINGS OF T H E A R N O L D A R B O R E T U M S TA F F J. H. Alexander 2002. Paraphyletic Syringa: evidence from sequences of nuclear ribosomal DNA ITS and ETS regions. Systematic Botany 27: 592?593 (with J. Li and D. Zhang). 2002. Phylogenetic relationships of Empetraceae inferred from sequences of gene matK and nuclear ribosomal DNA ITS region. Molecular Phylogenetics and Evolution 25: 306?315 (with J. Li et al.). P. Andersen 2002. The Institute for Cultural Landscape Studies of the Arnold Arboretum of Harvard University. In Restoring the Landscape: Policies for a New Sustainable Regional Project, ed. Francesca Leder. Ferrara, Italy: Facolta di Architettura di Ferrara. 2003. The Arnold Arboretum and the Early Years of Landscape Design Education in America. Arnoldia 62(3): 2?9. 2003. Spirit of Place (roundtable discussion). Architecture Boston 6(6): 8?17. 2003. Book review: Sacred Places: American Tourist Attractions in the Nineteenth Century, J. F. Sears. ArchitectureBoston, 6(6): 61. 2004. Book review: Becoming Cape Cod: Creating a Seaside Resort, J. C. O'Connell. ArchitectureBoston, 7(3): 49. J. L. Balzer 2007. Geographical distributions in tropical trees: can geographic range predict performance and habitat association in co-occurring tree species? Journal of Biogeography 34: 1916?1926 (with S. J. Davies et al.). 2007. Determinants of whole-plant light requirements in Bornean rain forest tree saplings. Journal of Ecology 95: 1205?1221 (with S. C. Thomas). 2007. Physiological and morphological correlates of whole-plant light compensation point in temperate deciduous tree seedlings. Oecologia 153: 209?223 (with S. C. Thomas). J. R. Bennett 2006. Phylogeny of the parasitic plant family Orobanchaceae inferred from phytochrome A. American Journal of Botany 93: 1039?1051 (with S. Mathews). D. E. Boufford 1992?2006. Harvard University Herbaria Databases: Botanists, Publications, Specimens. http:\/\/www.huh. harvard.edu\/databases (with K. N. Gandhi). 1998?2007. Biodiversity of the Hengduan Mountains Region, China. http:\/\/hengduan.huh.harvard.edu\/ fieldnotes (with Z. W. Ge et al.). 2002. Plant databases and the study of Asian Plants (1). Ouroboros 7(1): 10?13. 2002. Plant databases and the study of Asian Plants (2). Ouroboros 7(2): 13?15. 2003. Hengduan Mountains: International Hotspot of Biodiversity. In Xiuyuanjieying, ed. H. S. J. Wenji. Hong Kong: Commercial Press. 2003. A checklist of the vascular plants of Taiwan. In Flora of Taiwan, 2nd ed. Vol. 6. Taipai: National Taiwan University (with H. Ohashi et al.). 2003. Flora of Taiwan, 2nd ed. Vol. 6. Taipei: National Taiwan University (with C. F. Hsieh et al.). 2003. Rubus Linnaeus. In Flora of China, Vol. 9, eds. C. Y. Wu and P. H. Raven. Beijing: Science Press; St. Louis: Missouri Botanical Garden (with L. D. Lu). 2003. Phylogenetic position of Schnabelia, a genus endemic to China: evidence from sequences of cpDNA matK gene and nrDNA ITS regions. Chinese Science Bulletin 48(15): 1576?1580 (with S. H. Shi et al.). 2004. Mountains of Southwest China. In Hotspots Revisited: Earth's Biologically Richest and Most Endangered Ecoregions, 2nd ed., ed. R. A. Mittermeier et al. Mexico City: CEMEX Conservation International (with P. P. van Dijk). 41 2004. Japan. In Hotspots Revisited: Earth's Biologically Richest and Most Endangered Ecoregions, 2nd ed., ed. R. A. Mittermeier et al. Mexico City: CEMEX Conservation International. 2005. Cornaceae. In Flora of China, Vol. 8, ed. D. Y. Hong and P. H. Raven. Beijing: Science Press; St. Louis: Missouri Botanical Garden (with Q. Xiang). 2005. Aucubaceae. In Flora of China, Vol. 8, ed. D. Y. Hong and P. H. Raven. Beijing: Science Press; St. Louis: Missouri Botanical Garden (with Q. Xiang). 2005. Helwingiaceae. In Flora of China, Vol. 8, ed. D. Y. Hong and P. H. Raven. Beijing: Science Press; St. Louis: Missouri Botanical Garden (with Q. Xiang). 2005. Mastixiaceae. In Flora of China, Vol. 8, ed. D. Y. Hong and P. H. Raven. Beijing: Science Press; St. Louis: Missouri Botanical Garden (with Q. Xiang). 2005. Toricelliaceae. In Flora of China, Vol. 8, ed. D. Y. Hong and P. H. Raven. Beijing: Science Press; St. Louis: Missouri Botanical Garden (with Q. Xiang). 2005. Circaea lutetiana L. sensu lato (Onagraceae) reconsidered. Harvard Papers in Botany 9(2): 255?256. 2005. Polyploidy in the flora of the Hengduan Mountains Hotspot, Southwestern China. Annals of the Missouri Botanical Garden 92: 275?306 (with Z. L. Nie et al.). 2005. The Botanical Collections: Proceedings of the symposium Siebold in the 21st Century, Bulletin No. 41. Tokyo: University Museum, University of Tokyo (with H. Ohba). 2006. Angiospermae; Dicotyledonieae; Archichlamideae. Flora of Japan, Vol. IIa, ed. K. Iwatsuki et al. Tokyo: Kodansha, Ltd (with K. Iwatsuki and H. Ohba). 2007. Taxonomic note on Wikstroemia salicina (Thymelaeaceae). Acta Phytotaxonomica Sinica 45: 413?414 (with Y. H. Zhang and H. Sun). 2007. Taxonomic studies of Saussurea de Candolle (Asteraceae) in the Hengduan Mountains, China. An annotated list of recently collected specimens. Newsletter of Himalayan Botany 39: 13?37 (with K. D. Fujikawa et al.). 2007. Rhizophoraceae. In Flora of China, Vol. 13, ed. Z. Y. Wu et al. Beijing: Science Press; St. Louis: Missouri Botanical Garden (with H. N. Qin). 2007. Botanical Expedition to southern Gansu Province, China, May 2007. Newsletter of Himalayan Botany 40: 5?14 (with Y. Jia et al.). 2007. A Selection of Plants from Iriomote Island, Japan. Nantao, Taiwan: Endemic Species Research Institute and Research Center for Biodiversity, Academia Sinica (with C. Peng et al.). 2007. Meconopsis wilsonii subsp. wilsonii (Papaveraceae) rediscovered. Acta Botanica Yunnanica 29: 286?288 (with T. Yoshida and H. Sun). 2007. Atlas of the flora of New England: Cyperaceae. Rhodora 109: 237?360 (with R. A. Angelo). 2007. The Genera of Vascular Plants of Korea. Seoul: Academy Publishing Company (with C. Park, et al.). 2007. Two New Species of Wikstroemia (Thymelaeaceae) from Western Sichuan, China. Rhodora 109: 448? 455 (with Y. H. Zhang and H. Sun). Z. Chen 2003. Phylogeny of the Dipsacales s.l. based on chloroplast trnL-F and ndhF sequences. Molecular Phylogenetics and Evolution 26: 176?189 (with W. Zhang et al.). 2004. Regional differences in rates of plant speciation and molecular evolution: a comparison between eastern Asia and eastern North America. Evolution 58: 2175?2184 (with Q. Xiang et al.). 2004. Phylogenetics and biogeography of Alnus (Betulaceae) inferred from sequences of nuclear ribosomal DNA ITS region. International Journal of Plant Science 165: 325?335 (with J. Li). 2005. Phylogenetics of Betula (Betulaceae) inferred from sequences of nuclear ribosomal DNA. Rhodora 107: 69?86 (with J. Li and S. Shoup). 2007. Phylogenetic relationships of diploid species of Betula (Betulaceae) inferred from DNA sequences of nuclear nitrate reductase. Systematic Botany 32(2): 357?365 (with J. Li and S. Shoup). 2007. Phylogenetic and biogeographic diversification of Berberidaceae in the Northern Hemisphere. Systematic Botany 32(4): 731?742 (with W. Wang et al.). 42 2007. Mitochondrial matR sequences help to resolve deep phylogenetic relationships in rosids. BMC Evolutionary Biology 7: 217?231 (with X. Zhu et al.). S. Connor 2003. Shrubs and Vines at the Arnold Arboretum: A History. Arnoldia 62(2): 3?15. 2003. A picture is worth... The Public Garden 18(4): 39?41. 2004. In the Library: Hortus Nitidissimis. Arnoldia 63(1): 32. 2005. The Nature of Eastern Asia: Botanical and Cultural Images from the Arnold Arboretum Archives. Arnoldia 63(3): 34?44. R. E. Cook 2003. The Director's Report of the Arnold Arboretum: 1999?2002. Arnoldia 62(1). 2006. Botanical collections as a resource for research. The Public Garden 21(1): 18?21. 2007. The Future of Research at the Arnold Arboretum. Arnoldia 65(2) 23?29. J. Coop 2003. Sun-loving shrubs and vines for the Leventritt Garden. Arnoldia 62(2): 20?26 (with P. Del Tredici et al.). S. J. Davies 2002. The 52-Hectare Forest Dynamics Plot at Lambir Hills, Sarawak, Malaysia: Tree Distribution Maps, Diameter Tables, and Species Documentation. Diliman, Philippines: Center for Integrative and Development Studies, University of the Philippines (with H. S. Lee et al.). 2003. Effects of nutrient addition, mulching and planting-hole size on early performance of Dryobalanops aromatica and Shorea parvifolia in enrichment plantings in Sarawak, Malaysia. Forest Ecology and Management 180: 261?271 (with A. Vincent). 2003. Coadaptation and coevolution of Macaranga trees and their symbiotic ants. In Genes, Behaviours and Evolution of Social Insects, ed. T. Kikuchi et al. Sapporo, Japan: Hokkaido University Press (with T. Itino and T. Itioka). 2003. The trees of Pasoh Forest: stand structure and floristic composition of the 50-hectare forest research plot. In Pasoh: Ecology and natural history of a Southeast Asian lowland tropical rain forest, ed. T. Okuda et al. Tokyo, Japan: Springer (with M. N. Nur Supardi et al.). 2003. Lambir forest dynamics plot, Sarawak, Malaysia. In Forest Diversity and Dyna","distinct_key":"arnoldia-2008-Director's Report: 2003-2008"},{"has_event_date":0,"type":"arnoldia","title":"Director's Report: 2003-2008","article_sequence":1,"start_page":1,"end_page":53,"url":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/action\/directLinkImage?assetId=25437","featured_photo":"https:\/\/assetbank.arboretum.harvard.edu\/assetbank-aahu\/servlet\/display?file=b34509f9d4eed170b36b.jpg","volume":65,"issue_number":4,"year":2008,"series":null,"season":null,"authors":"Schulhof, Richard; Cook, Robert E.","article_content":"mism, ed. E. Losos and E. Leigh. Chicago: The University of Chicago Press (with H. S. Lee et al.). 2003. Book review: A Manual for Forest Plantation Establishment in Malaysia, ed. B. Krishnapillay. Journal of Tropical Forest Science 15(2): 365?367. 2004. Habitat heterogeneity and niche structure of trees in two tropical rain forests. Oecologia 139: 446?453 (with M. D. Potts et al.). 2004. Soil-related habitat specialization in dipterocarp rain forest tree species in Borneo. Journal of Ecology 92: 609?623 (with P. Palmiotto et al.). 2004. Evolution of host affiliation in Crematogaster (Formicidae) inhabitants of Macaranga (Euphorbiaceae). Evolution 58: 554?570 (with S. P. Quek et al.). 2004. Lambir Hills Forest Dynamic Plot, Sarawak, Malaysia. In Forest Diversity and Dynamism: Findings from a Network of Large-Scale Tropical Forest Plots, ed. E. Losos and E. G. Leigh. Chicago: University of Chicago Press (with H. S. Lee et al.). 2004. Sinharaja 25-hectare plot: comparisons with other forests in the CTFS network. In Ecology of Sinharaja Rain Forest and the Forest Dynamics Plot in Sri Lanka's Natural World Heritage Site, ed. C. V. S. Gunatilleke et al. Columbo, Sri Lanka: WHT Publications. 2004. Palanan forest dynamics plot, Philippines. In Forest Diversity and Dynamism, ed. E. Losos and E. G. Leigh. Chicago: The University of Chicago Press (with L. L. Co et al.). 2005. Tropical tree -diversity: results from a worldwide network of large plots. Biologiske Skrifter 55: 565? 582 (with R. Condit et al.). 43 2005. Tree growth is related to light interception and wood density in two mixed dipterocarp forests of Malaysia. Functional Ecology 19: 445?453 (with D. A. King et al.). 2005. Edaphically-associated variation in performance correlates with species' distribution patterns in a Bornean rainforest. Journal of Ecology 93: 879?889 (with S. E. Russo et al.). 2005. Forest Trees of Bukit Timah, Singapore: Population Ecology in a Forest Fragment. Singapore: Simply Green (with J. V. LaFrankie et al.). 2005. Soil-related floristic variation in a hyperdiverse dipterocarp forest. In Pollination Ecology and the Rain Forest (Sarawak Studies), ed. D. Roubik et al. New York: Springer Science (with S. Tan et al.). 2006. Regeneration of native plant species in restored forests on degraded lands in Singapore. Forest Ecology and Management 237: 574?582 (with K. Shono and Y. K. Chua). 2006. Spatial associations of humus, nutrients, and soils in mixed dipterocarp forest at Lambir, Sarawak, Malaysian Borneo. Journal of Tropical Ecology 22: 543?553 (with I. C. Baillie et al.). 2006. The role of wood density and stem support costs in the growth and mortality of tropical trees. Journal of Ecology 94: 670?680 (with D. A. King et al.). 2006. The importance of demographic niches to tree diversity. Science 313: 98?101 (with R. Condit et al.). 2006. Competitive dominance in a secondary successional rain forest community in Borneo. Journal of Tropical Ecology 22: 53?64 (with H. Semui). 2006. Contrasting structure and composition of the understory in species-rich tropical rain forests. Ecology 87(9): 2298?2305 (with J. V. LaFrankie et al.). 2006. Comparing tropical forest tree size distributions with the predictions of metabolic ecology and equilibrium models. Ecology Letters 9: 589?602 (with H. C. Muller-Landau et al.). 2006. Testing metabolic ecology theory for allometric scaling of tree size, growth, and mortality in tropical forests. Ecology Letters 9: 575?588 (with H. C. Muller-Landau et al.). 2006. Nonrandom processes maintain diversity in tropical forests. Science 311: 527?531 (with C. Wills et al.). 2006. Growth and mortality are related to adult tree size in a Malaysian mixed dipterocarp forest. Forest Ecology and Management 223: 152?158 (with D. A. King et al.). 2006. Forest Trees of Palanan, Philippines: A Study in Population Ecology. Diliman, Philippines: Center for Integrative and Development Studies, University of the Philippines (with L. L. Co et al.). 2007. The role of gap-phase processes in the long-term biomass dynamics of four old-growth tropical forests. 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