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Department of Botany & the U.S. National Herbarium TheThe PlantPlant PressPress

New Series - Vol. 8 - No. 1 January-March 2005 Botany Profile 150 Years of Thistles, Daisies, and Sunflowers By Vicki A. Funk he US National Herbarium has an from forests to high elevation grasslands, of the herbarium but mostly during the active research and collecting however they are less common in tropical last 100 years. A series of synanther- Tprogram in the flowering wet forests and more common in open ologists (see photos on Page 10) have family Compositae (). Recent areas. enhanced the herbarium by their accomplishments include the Helianthe- The general perception of this family collecting efforts and their willingness ae treatment for Flora of Ecuador, a as “weedy” is only partially correct. to identify Compositae from throughout supertree for the entire family, a new Certainly there are members that benefit the world. I think it is safe to say that international organization and website, from disturbance but most species have a anyone wanting to work in Neotropical and the near completion of the mono- restricted distribution, and just about Compositae must visit or borrow the graph on the Espeletiinae by José every “at risk” habitat in the world collections at US. Cuatrecasas. contains members of this family that are The first people who made major The Compositae (Asteraceae) an important part of the flora. In fact, the contributions to the Compositae contains the largest number of described 1997 IUCN Red List of Threatened collection were Asa Gray and Benjamin species of any plant family, 24,000— (Walter and Gillett 1997) has L. Robinson, both synantherologists with estimates of total species ranging 2,553 species of Compositae on its list. from Harvard. Gray identified the plants up to 35,000—distributed in 1,600 – It is an important family not only housed in the U.S. National Herbarium 2,000 genera occurring on all continents because it is pervasive in many ecosys- that were collected during early U.S. except Antarctica. Estimates vary, but tems but also because members of the expeditions, Plantae wrightianae texano- assuming that there are 200,000 – family dominate our gardens (about 260 neo-mexicanae, the boundary survey, 300,000 species of flowering plants, species in the U.S.), provide a number of published in Smithsonian Contributions then one out of every 8 – 12 species of invasive species (i.e., Chromolena, Cirsi- to Knowledge 3, Art. 5. 1852 and 5 (Art flowering plants is in the Compositae um, Taraxicum), produce secondary plant 6). 1853 and the United States Explor- (about 10 percent). The family is products that are used in traditional and ing Expedition under the command of monophyletic, characterized by florets homeopathic medicines (ca. 500 in China Charles Wilkes, published 1854 and arranged on a in centripetal alone) and in pesticides, and provide food 1856. Likewise, Robinson identified heads and surrounded by , by (i.e., dandelions, sunflowers, artichokes, many specimens from US including the anthers fused in a ring with the lettuce). material from Hitchcock, Killip and pushed or brushed out by the style, and Smith, Pringle, A.C. Smith, and Rose. by the presence of (cypselas) he US National Herbarium houses Sidney Fay Blake, a student of often with a pappus (see figure on Page approximately 500,000 specimens Robinson, was one of the foremost 9). Although the family is well-defined, Tof Compositae and 8,500 recorded Compositae workers of the 20th century. there is a great deal of variation among types and although it has global represen- In between his master’s and his Ph.D. he the members: the habit varies from tation it is strongest in the Neotropical was perhaps one of our first “interns” annual and perennial herbs to shrubs, Compositae, perhaps the best in the because he spent the summer of 1913 vines, or trees, although few are true world. There are no doubt many speci- arranging plants at the Smithsonian. epiphytes; the heads can have one to mens in the general herbarium that should Upon the completion of his Ph.D. Blake more than 1,000 florets; chromosome be in the type collection but have not yet was offered jobs at both the Smithsoni- numbers range from n = 2 to high level been identified as such or they are an and the Department of Agriculture. polyploidy with n = 114; and species duplicate material that was, until recently, He took the USDA job because the pay grow in just about every type of habitat kept in the main herbarium. This large collection was assembled over the history Continued on page 9 Visitors

Michael Nee, New York Botanical German Carnevali, Independent David Rhode, Desert Research Institute, Garden; Solanaceae and Cucurbitaceae researcher; Flora de Guaramacal (Orchid- Neveda; Death Valley Expedition (Coville (10/4). aceae) (10/27-11/3). & Funston) (11/16-11/17). Marisol Toledo, Missouri Botanical Richard Clinebell, Missouri Botanical Birgit Gemeinholzer, Botanischer Garden; Bolivian Meliaceae (10/4). Garden; Onagraceae pollination evolution Garten und Botanisches Museum Berlin- (11/2-11/6). Dahlem, Germany; Phylogeny of Tribe Janice Dern, Independent researcher; Lactuceae (Compositae) (11/19-11/22). volunteer interview (10/5). Ralph and Pat Dixon, Legacy Society of Smithsonian Institution; Present gift of Robert Bye, Universidad Nacional Patricia Gomez Bustamante, Indepen- 1924 scrapbook of wildflowers along Autónoma de México, Instituto de dent researcher; volunteer interview (10/ C&O Towpath (11/2). Biologia; Flora of northern Mexico and 5). Edward Palmer collections (11/22). Peter Hoch, Missouri Botanical Garden; H. David Clarke, University of North Onagraceae (11/2-11/6). Alison Colwell, US Geological Survey, Carolina, Asheville; Identification of Western Fisheries Research Center; Guyana plant collections (10/6-10/9). Rachel Levin, Amherst College; Onagra- Orobanche, Platanthera, Plagiobothrys ceae (11/2-11/6). Chad Hasby, Florida International from Yosemite National Park (11/22). University; Equisetaceae (10/14). Robert Raguso, University of South Roger Troutman, Independent re- Carolina at Columbia; Onagraceae searcher; Liatris (Asteraceae) E.S. Steele Steve Glenn and Gerry Moore, Brook- pollinator systems (11/4-11/7). lyn Botanical Garden; Woody plants for collections (11/22-12/13). New York Metropolitan Project (10/19-10/21). Madison; Onagraceae (11/4-11/6). Católica del Ecuador; Trees of Yasuni Richard Olmstead, University of Wash- Steve Junak, Santa Barbara Botanic Gar- Forest Dynamics Project, Ecuador (11/ ington; Morphology of Lamiales (10/19- den; Flora of Channel Islands, California 29-12/27). 4/20). (11/16-11/18). Continued on page 8 L. Alan Prather, Michigan State Univer- sity; Lamiaceae and Polemoniaceae. (10/ Travel 26-10/29). Mark Littler, Diane Littler and for Botanical Documentation; and to Barrett Brooks traveled to Panama (9/ Kauai, Hawaii (11/18 – 12/4) to work on 14 – 9/30) to continue their work on the the Flora of the Marquesas Islands. “Marine Flora of the Caribbean and W. John Kress and Mike Bordelon The Plant Press Pacific Panama” project. traveled to Yangon, Myanmar (11/3 – 11/ New Series - Vol. 8 - No. 1 Rusty Russell traveled to the Univer- 20) to collect Zingiberales and other sity of California Reserve-Deep Canyon flowering plants. Chair of Botany outside Palm Desert, California (9/23 – Lawrence Skog traveled to Toronto, W. John Kress 9/27) to meet with colleagues from the Canada (11/12 – 11/15) to present an ([email protected]) Bureau of Land Management, University invited lecture, “Looking Back at Gesner- of California Riverside, University of ia,” to the Toronto Gesneriad Society and EDITORIAL STAFF Redlands and Riverside Municipal to visit the Botany Department at the Editor Museum on developing GIS tools that University of Toronto; and to Philadel- Gary Krupnick incorporate historical specimen data; and phia, Pennsylvania (11/21 – 11/24) to ([email protected]) to Seattle, Washington (10/31 – 11/2) to study and record specimens of Gesneria- present a special symposium, “The ceae at the herbarium of the Academy of News Contacts Botanical Legacy of the Lewis & Clark Natural Sciences. MaryAnn Apicelli, Robert Faden, Ellen Expedition” at the annual meeting of the Maria Faust traveled to Florida Keys, Farr, George Russell, Alice Tangerini, and Elizabeth Zimmer Crop Science Society of America. Florida (11/14 – 11/18) to collect Paula DePriest traveled to Raleigh, harmful dinoflagellates. The Plant Press is a quarterly publication provided North Carolina (10/21 – 10/22) to Dan Nicolson traveled to Kuching, free of charge. If you would like to be added to the participate in the Ph.D. defense of Sarawak in Malaysia (11/19 – 11/27) to mailing list, please contact Gary Krupnick at: th Department of Botany, Smithsonian Institution, PO Rebecca Yahr at Duke University. attend the 9 International Aroid Sympo- Box 37012, NMNH MRC-166, Washington, DC Alice Tangerini traveled to Pitts- sium; and to the Royal Botanic Gardens, 20013-7012, or by E-mail: [email protected]. burgh, Pennsylvania (10/21 – 10/24) to Kew in London, U.K. (11/27 – 12/4) to Web site: http://www.nmnh.si.edu/botany attend a meeting of the American Society work on the Taxonomic Literature, of Botanical Artists at the Hunt Institute edition 2 (TL-2).

Page 2 Plants Will Talk with Plant Talk Chair he Convention on Biological Diversity (CBD) standing and documenting plant diversity; 2) conserv- Chair initiated at the Earth Summit in Rio de Janeiro ing plant diversity; 3) using plant diversity sustain- Tin 1992 focused the nations of the world on the ably; 4) promoting education and awareness about environmental crisis affecting the entire planet. The plant diversity; and 5) building capacity for the With treaty set the stage for an international effort to conservation of plant diversity. Within these broad document, use, and conserve biodiversity, with many categories are 16 specific targets to be achieved in countries implementing rules and regulations directed the next five years, such as, “Target 4: at least 10% of at their national natural heritage. However, it was not each of the world’s ecological regions effectively A until ten years after Rio in 2002 that a major step was conserved;” “Target 7: 60% of the world’s threatened taken to specifically protect the world’s plant species. species conserved in situ;” “Target 9: 70% of the Following a series of discussions, meetings, and genetic diversity of crops and other major socio- View declarations by major botanical and biodiversity economicaly valuable plant species conserved, and institutions, such as Botanic Gardens Conservation associated indigenous and local knowledge main- International, Royal Botanic Gardens at Kew and tained;” and “Target 12: 30% of plant-based products Edinburgh, Missouri Botanical Gardens, IUCN derived form sources that are sustainably managed.” Species Survival Commission, Instituto Alexander von These goals are not trivial and will take international W. Humboldt, and the Smithsonian among many others, efforts to achieve in the next five years. the Global Strategy for Plant Conservation (GSPC; As natural history museum scientists, we can take John see http://www.biodiv.org/decisions/) was officially part in furthering each of these targets. Perhaps Kress adopted during the 6th Conference of the Parties in Target 1, to produce “a widely accessible working list The Hague, Netherlands. This action was a monumen- of known plant species, as a step towards a complete tal achievement for a number of important reasons. world flora,” is the goal to which we can best contrib- First of all, the Global Strategy for Plant Conserva- ute. The Department of Botany, however, has recently tion put the Earth’s plants at the forefront of conserva- taken a large step in helping achieve all of the targets tion efforts. Previously “panda-envy” was common of the Global Strategy by forming a partnership with among plant conservationists who were unsuccessfully four other plant-based organizations to jointly spon- trying to interest the general public in setting the same sor and promote the international magazine Plant priorities and providing the same level of protection Talk, a publication dedicated to advancing in situ for photosynthetic species as they were for various plant conservation (see http://www.plant-talk.org). large and furry or feathered animals. The basic Together with The Eden Project (based in Cornwall, ecological function that plants provide as the corner- UK), the Royal Botanic Gardens at Kew, the EU- stone of both terrestrial and many marine ecosystems based Plantlife International, the Critical Ecosystem was being overlooked in favor of the charismatic and Partnership Fund (part of Conservation International), endangered large mammals and birds. The GSPC and the Artdatabanken (the Swedish Species Informa- changed that perspective by providing the first global tion Center), Botany at the Smithsonian, in collabora- conservation strategy for a specific group of organ- tion with our partner on the Mall, the United States isms recognized by the Convention on Biological Botanic Garden, have joined forces to transform the Diversity. ten-year old Plant Talk into a widely distributed Second, and most important, the Global Strategy magazine that will seek to advance all 16 targets of for Plant Conservation is a pioneering plan adopted by the GSPC through news, views, and reports on plant the CBD that provides concrete targets and activities conservation around the world. that are to be achieved within a given period of time. The new Plant Talk consortium will work hand-in- Although the CBD had previously set up a broad hand with our own Plant Conservation Unit as well as framework for using and conserving biodiversity, the scientists in the Department of Botany at the Smith- GSPC is the first international plan that includes sonian to understand and conserve the immense plant concrete targets for the sustainable use and protection diversity of our planet. of the world’s plants by a specific date: the year 2010. The strategy and targets put forward by the group of plant conservation activists that crafted the GSPC was radical enough, but the most startling event was that the strategy was actually embraced and adopted by the CBD Conference of the Parties. What is in the Global Strategy that makes it so significant? The plan is divided into five sections with specific targets listed under each section: 1) under-

Page 3 David Lorence, curator at NTBG, Washington, DC area. Bob Faden was Staff Tangerini was able to illustrate, in pencil, chairman of the chapter for four years eleven species with nine of them being and chaired the 2002 Eastern Winter Research & described as new. Although the majority Study Weekend, which the local chapter Activities of the plants were dried specimens, organized on behalf of NARGS. Audrey Wagner and Lorence provided excellent Faden has been involved in the chapter in color images of living material to aid in many capacities, including organizing the On 3 November, W. John Kress and the reconstruction. The staff was friendly plant propagation and sales during the Mike Bordelon traveled to Yangon, and helpful; Tim Flynn looked for addi- 2002 meeting. She has volunteered in the Myanmar to collect Zingiberales and tional plant material and provided work Botany Research Greenhouse for more other flowering plants. While in Myan- space and any small tools that Tangerini than 24 years where she helps maintain mar, Kress presented a training workshop needed for dissecting; the NTBG front the Commelinaceae collections. on pollination biology and plant office helped with any photocopying to a group of students at the University of problems that arose and were patient as W. John Kress and colleagues Jiang Yun Yangon. On 18 November, returning via Tangerini spread out the numerous Gao and Qing-Jun Li, both at Xishaung- Bangkok, Thailand, Kress and Bordelon unmounted collections all over the floor banna Tropical Botanical Garden in attended the 3rd IUCN World Conserva- of the photocopy room. Tangerini was Yunnan, China, have received a two-year tion Congress and assist with a Smith- introduced to two famous plant collec- $25,000 grant from the Chinese Natural sonian Institution poster presentation by tors: Ken Wood and Steve Perlman, who Science Foundation for their project Leonard Hirsch, Senior Policy Advisor. provided many of the specimens. The “Research on the Effects of Floral grounds of NTBG were extensive and even Longevity on Male and Female Fitness From 19 November to 4 December, Dan after a tour given by Lorence late one and its Evolutionary Implications in Nicolson traveled to Kuching, Sarawak in afternoon there was just not enough time Hedychium (Zingiberaceae).” Members Malaysia and the Royal Botanic Gardens, to even see it all. of Hedychium display many different Kew in London, U.K. In Kuching, floral characteristics from other genera Nicolson attended the 9th International Alain Touwaide delivered a paper, in the Zingiberaceae, such as conspicuous Aroid Symposium, where he chaired two “Medicine Across Cultures: 600-1600,” longer floral longevity, asymmetrical sessions and presented the final lecture, at the 19th Barnard Medieval and Renais- flower and unique structure of sexual “Aroid Seekers in the Great Forests of sance Conference at Barnard College, flowering organs. These unique floral Malesia.” On the final day, Nicolson Columbia University, on 4 December. characteristics indicate that Hedychium spoke to botany students at the University Touwaide delivered another paper, “The plants have a rather unique pollination of Malaysia (Unimas), where he talked Greeks and the Irrational Revisited,” at mechanism and breeding system. The about the “Roots of a Botanist,” as an Johns Hopkins University in Baltimore, project is designed to conduct research encouragement for them to find their Maryland, on 7 December. on floral biology, pollination ecology and passion. Nicolson also had the opportu- mating system of four Hedychium nity to climb Matang Hill where he first Warren Wagner gave a talk entitled “The species and varieties, which show collected Schismatoglottis nicolsonii Surprising Fate of Hawaiian Lineages” at different floral longevity and are all over 40 years ago. On his return trip, he the National Tropical Botanical Garden distributed in South Yunnan, China. The stopped for a week at the Royal Botanic for the ‘Science in the Garden’ series in research also includes a series of Gardens, Kew, to work the Taxonomic Kalaheo, Kauai on September 29, and a manipulation experiments to compare the Literature edition 2 (TL-2), checking talk entitled “Understanding Extinction in effects of floral longevity on male and their reprint files. Hawaii” for the Kauai Native Plant female fitness. Society in Lihue, Kauai on November 20. On October 29, Paul Peterson gave an invited lecture at the United States Botanic Garden entitled “Grasses and Other Plants of the Sierra Tarahumara.” The research of Peterson and other Botany staff is featured in the current USBG exhibit “Going Where They Grow: Awards & Exploring the Front Lines of Botany.” Grants On 18 November Alice Tangerini traveled to the island of Kauai to work on Botany curator Robert Faden and his the Flora of the Marquesas Islands as part wife, Smithsonian Behind the Scenes of a grant secured by Warren Wagner. Volunteer Audrey Faden, were given an Illustrating in the setting of the beautiful Award of Merit by the North American National Tropical Botanical Garden was Rock Garden Society (NARGS) for inspiring and productive. Under the service to the local Potomac Valley Hedychium bordelonianum from direction of Wagner and collaborator, Chapter of NARGS, which is based in the Southeast Asia

Page 4 reefs, sea grass beds, sand, and various Study of Dinoflagellates Connected to species of macroalgae. Using a combina- Harmful Toxins tion of species taxonomy and genetic Marine dinoflagellates (also known as in Belize to study the ecological mor- sequencing techniques, molecular assays phytoplankton) are microscopic, single phology and taxonomy of dinoflagellates. are being developed to advance the study celled organisms that drift with the ocean Using medical molecular biology applica- of the ecology, biodiversity, and distribu- currents and are responsible for the tions, scanning electron microscopy tion of these toxin producing organisms. majority of the flux of organic matter to (SEM) and other cutting edge techniques, both higher tropic levels and the ocean they hope to establish a foundation for interior. The community structure and understanding the biodiversity of tropical ecological function of contemporary coastal waters. marine ecosystems are critically depen- During November, Maria A. Faust dent upon these organisms. Their mor- joined Karen A. Steidinger, FIO-Florida phology and ecology are virtually un- Marine Research Institute, St. Petersburg, known. Fl; Mark W. Vandersea and William Many dinoflagellate species synthe- Holland, Center for Coastal Fisheries and size harmful toxins and are responsible Habitat Research, NOAA, Beaufort, NC; for ciguatera fish poisoning which affects and other members of the Harmful Algal higher life forms in marine environments. Bloom Project to collect tropical Worldwide ciguatera poisoning causes dinoflagellates in the Florida Keys that more death and illness to humans than all produce ciguatoxins to better understand other harmful algal toxins combined. their ecology, biodiversity, and toxicol- Smithsonian scientists have been ogy. Samples yielded rich flora of dino- SEM image of Gambierdiscus toxicus, conducting ongoing research within the flagellates representing great topological a ciguatoxin-producing dinoflagellate coral reef – mangroves at Carrie Bow Cay diversity in the phytoplankton, patch [100 um long x 80 um wide] (Adachi and Fukuyo 1979).

15-16 April 2005 NATIONAL MUSEUM OF NATURAL HISTORY In collaboration with the United States Botanic Garden Sponsored by the International Association for Plant Taxonomy “The Future of Floras: New Frameworks, New Technologies, New Uses” For centuries botanists have created regional floras for the purposes of inventory, identification, description, and classification. But what will the floras of the future look like? Will there be a need for floras when a Web-based “Encyclopedia of Life” becomes a reality? What will field taxonomists be doing in 20 years? New technologies are now being developed to facilitate the coupling of field work with ready access of plant data that exist in biodiversity institutions. The development of electronic field guides and image identification systems, as well as DNA barcoding methods to identify species in the field, have great potential to augment if not completely replace the traditional paper-based flora. Some scientists are concerned that new technologies that further the inventory and classification of life may also threaten the field of taxonomy. Will new technologies replace taxonomists who work directly with specimens? Will new techniques provide faulty identifications? Proponents of the new technologies believe that the easier it is for end-users to employ good taxonomic data for identification, the more taxono- mists will be appreciated for their skills and knowledge. As the debate continues, floras built on Web-, image-, and DNA-based products are fulfilling new functions that paper-based floras have not been able to attain. All of these topics focusing on “Vernonia salviniæ, Hemsl.” (Tab. 41) the floras of the future will be discussed and debated at the symposium. in Biologia Centrali-Americana, Botany, Vol. V (1879-1888) Information and registration at: http://persoon.si.edu/sbs/ by William B. Hemsley Fax: 202-786-2563 – e-mail: [email protected] [Image Courtesy Smithsonian Institution Libraries]

Page 5 the United States Botanic Garden for not far from the west end of New New Species of educationl purposes. Entandrophragma Guinea. Mortoniodendron is angolense (Meliaceae), an African “This photograph will be placed mahogany, was grown from a seed with the specimen in our herbarium, Described collected around the stump of an illegally for Prof. Buswell kindly gave us the Laurence Dorr and his colleague, cut tree on the slopes of Étinde (Little latter.” Tom Wendt from the University of Texas Mount Cameroon), Cameroon in 1986. Seeing all this, Dan Nicolson checked at Austin, describe the taxonomic history Seeds of Psychotria riparia (Rubiaceae), the herbarium and, sure enough, Egbert of the genus Mortoniodendron in the a red-fruited shrub or small tree in one of Walker (former curator of Old World latest issue of Lundellia (7: 44-52. the largest genera of the coffee family, botany) had mounted the photograph 2004). The genus was named in honor of were collected in the Taita Hills, Kenya in (Neg. 37411-C), annotated the black and Conrad V. Morton (1905-1972) who 1985 when Faden participated in the white photo with color notes. Nicolson served for many years as a curator in National Museums of Kenya Taita Hills found that the Smithsonian also had only Botany. Characters that define M. penta- Expedition. one specimen of the species, a collection gonum, a species known from a limited made in Amboina in 1913 by C.B. Robin- area in Guatemala, are elaborated in the son, who was killed there, but that is paper. A closely related, but distinct another story. species from Veracruz, Mexico, M. uxpa- napense Dorr & T. Wendt, is described for the first time and is accompanied by Touwaide and illustrations by Alice Tangerini. The new Appetiti Join species is considered to be Critically Endangered (CR) under the IUCN Red Botany List classification for endangered Botany Receives Alain Touwaide and Emanuela Ap- species. petiti have joined Botany as new tempo- Historic rary trust fund staff funded by a four-year Watercolor grant from the National Institutes of Health. They have both been in Botany for By Dan H. Nicolson a number of years as volunteer Research On 18 November, Frederick ‘Ted’ Associates working with Dan Nicolson Bayer, Curator of Invertebrate Zoology and W. John Kress on ancient botany and for more than 40 years, made a gift to medicinal plants. The grant will provide Botany—a small, original watercolor, the resources to allow them to continue annotated in pencil, “Bauhinia: Yellow their work on a computerized database of Ebony Vine, Biak, near Sarido - 11 April medicinal plants used between the 5th [19]45.” Bayer was then stationed on century B.C. and the 2nd century A.D., a Biak, an island off the back of the “bird’s priority project recognized by the NIH. head” of New Guinea. The relevant texts, many of which are in The gift included two letters. One was Touwaide’s library, will be read in the dated 3 June 1945 from W. M. Buswell original Greek and translated into English (Coral Gables, Florida), which said, “I for entry into the database. received the Bauhinia specimen but haven’t been able to name it yet.” The other letter, dated 27 July 1946, is to Bayer (then living in Florida) from E. D. Merrill (then at the Arnold Arboretum): “Dr. Walker of the Smithsonian Cover of Lundellia with photograph of Institution has sent me a very nice Mortoniodendron uxpanapense. photographic print of your sketch of the New Guinea plant [that] I deter- Greenhouse mined for Prof. Buswell as Bauhinia lingua DC. The latter was described Donates African from specimens collected on Amboina Island, and we have one collection of it Plants to USBG from Amboina; yours was the first The Botany Research Greenhouse record from New Guinea, and the first donated two African woody plants grown record of the species from outside of from seed collected by Robert Faden to the small island of Amboina which is

Page 6 pollination, evidenced by red tubular source for dispersing birds. These shifts Speculations about flowers that differ from those of their underscore the reciprocal relationship Speciation presumed ancestor. More dramatically, in between evolution and ecology and four lineages a capsular fruit has evolved caution against drawing too many conclu- Why do some plants diversify through into a fleshy fruit that is used as a food sions about cause and effect. the process of speciation? Jonathan Price and Warren Wagner attempt to answer this question in a study published in the October issue of Evolution, “Speciation in Hawaiian angiosperm lineages: cause, consequence, and mode” (58: 2185-2200. 2004). The Hawaiian flora provides an ideal opportunity to pose questions about speciation thanks to its extreme isolation, self-contained geologic history and highly unique flora. All Hawaiian plant species are descended from a limited and identifiable set of original colonists. Some of these, such as Various members of the Hawaiian silversword alliance. the famous Silversword alliance, diversi- fied into many distinct species, while Correction values of the flowering cherry—seen as others remained a single species. By a sign of civilization—they planted their comparing such features as how species In the 2004 special Symposium Issue of the Plant Press (Vol. 7 – No. 3), an own cherry in the imperial palace in are pollinated and dispersed, and what Kyoto in 1357, replacing the Mountain range of islands and habitats they occupy, inaccurate summary of a talk by Wybe Kuitert was included. To provide a better Cherry. This new imperial cherry was a Price and Wagner were able test weather form of the Oshima Cherry (P. serrulate certain ecological characters drive the description of his paper, Kuitert’s abstract is as follows: var. speciosa), native to regions about diversification process. 400 kilometers to the east of Kyoto, Dispersal appeared to be the most Wybe Kuitert proving that cherries were transported important feature in speciation. Price and Kyoto University of Art and Design over long distances. Hitherto, isolated Wagner believe on the one hand the “Political Change and Cultural Values of plant material of the highly variable lineages with poor dispersal ability (those Oshima Cherry was brought to Kyoto with large seeds, or those that depend on Plants: Origins of Cherry Hybridization in Medieval Japan” from the east, while the Mountain floatation) had few species because they Cherry had been brought to the city a tended not to colonize very many islands Humans played a dramatic role in the century earlier from Yoshino in the and thus could not readily establish evolution of Japan’s flowering cherries. south. Cherry populations that had isolated populations that begin the In the 5th century, a wave of immigrants developed separate identities over the speciation process. On the other hand, arrived from the continent. Accommo- course of their evolution were brought lineages with highly dispersible fruits dating the new population meant clearing together and could start to hybridize, a (especially those sought out and ingested forests, which gave new opportunities process that would lead to a number of by birds) also have on average few species for light-loving plants, like flowering distinctly different garden forms in the because they colonize many different cherries, to flourish. In 834 AD, the succeeding centuries. islands but disperse too much permitting plum tree (Prunus mume), traditionally From the end of the 17th century, gene flow between populations. Those planted with citrus trees in front of the cherries have been recorded in the that are dispersed by external adhesion to imperial palace in the capital city written history of Ninna-ji, still famous birds (with barbs or sticky surfaces, or (present Kyoto), was replaced by the for its classic garden forms and the with tiny seeds that become embedded on Mountain Cherry (P. serrulata var. Hirano Shrine. The region of Ninna-ji bird’s feet) had more species than those spontanea Lindley). This tree was and Hirano Shrine, in between Arashiya- with other modes of dispersal. This may brought from the mountainsides of ma, with its Japanese Mountain Cherries be because they disperse enough to Yoshino, close to Nara, an earlier capital from Yoshino and the center of the city, colonize all major islands, but not so city. Over the course of the 13th century, must have been an increasingly interest- much that populations continue to large numbers of cherries were brought ing cherry-hybrid area beginning in the interbreed. from Yoshino, and planted at Arashiyama 14th century when Oshima Cherries were Despite all of this, in many cases and in the outskirts of Kyoto. planted in the imperial palace. Their especially species-rich groups, the type With changing political winds, hybrids, the historic garden forms of of dispersal or pollination shifted, often warlords from eastern Japan occupied Ninna-ji and Hirano Shrine are indeed through the evolution of unique, special- the capital in 1336 and installed their spectacular garden plants that have stood ized features. For example, in eleven own emperor. Well-aware of the cultural the ages. lineages there was a shift toward bird

Page 7 The Heart of the Herbarium By Heijia L. Wheeler, Volunteer In a brightly lit room, two volunteers quietly study the pressed plant specimens they are about to mount. They are two of the eleven volunteers who use their skills and artistic senses to help Katherine “Kat” Rankin take the dried specimens that arrive in their plebian newspaper wrappings and turn them into works of art. These beautifully mounted specimens make up the heart of the U.S. National Herbarium. Even as researchers are using current technologies such as DNA bar- coding, electronic field guides, and digital imaging, they still rely on the physical plant sample for detailed study of glands, hairs, cellular structures, anatomy and chemical analysis. It would be futile to toss a digital image or a photographic slide into a test tube for a chemical analysis of floral fragrances, no matter how beautiful the image. The environment in which these volunteers work is organized and quiet, away from their other worlds of responsi- Volunteer Larry Owens mounting a specimen at the US National Herbarium. bility. The volunteers are aware of the (Photo by Deborah Bell) value of their contribution and they are a part of the family of Botany and the The Herbarium has over 4.7 million resulting mounted specimens are estheti- Smithsonian Institution. It is little wonder specimens of which 96,000 are type cally pleasing. This is a synergistic that so many of them have been volunteer- specimens of all plant groups. Every day relationship. The Herbarium’s gain is ing for so long. Mary Starr, Mary Ellen new specimens arrive at the Herbarium obvious. Last year the volunteers mounted Wiser, and Gwen Petitjean have been from collectors all over the world. These over 3,500 specimens. The volunteers in volunteering since the mid 1980s. They are donated by scientists from Alabama to turn gained a sense of accomplishment have a combined total of 60 years of Australia. The researchers at the and the pleasure of each others company. volunteer service to this project. Marga- Smithsonian also bring back specimens ret Schweitzer has been a volunteer for 15 related to their research projects from the Visitors world over. All of these specimens are years and the newer volunteers, Joan Lorr, Continued from page 2 Jo Ann McCann, Larry Owens, Pat carefully dried, recorded and sent to Rankin’s group who turn them into Trisdorfer, Cate Puzo, Anne Datko, and Curt Pueschel, State University of New Silvia Domenge are just as loyal and permanent vouchers to be placed in their proper spot in the Herbarium. York at Binghamton; Algal Collection dedicated. They range from retirees as old (12/2). as 87 years to working individuals as The mounting process is very precise. young as 45 years, all of them determined They use the strapping and stitching Ihsan Al-Shehbaz, Missouri Botanical to make a contribution. The proof is in the method rather than simply gluing the plant Garden; Arabis (Brassicaceae) Flora years of loyal hard work they have given, to the mounting paper. The U.S. National North America (12/6-12/8). and the thousands of specimens they have Herbarium is one of the few herbaria that USDA Graduate School class, U.S. mounted. One gentleman was so adamant use this method for plant mounting. All Department of Agriculture Graduate to remain a volunteer when he had to the mounters get fundamental instructions School, Woody Plant Identification Class; move away from the DC area last year that and rules in their training. Once they are Herbarium tour (12/6; 12/20). he continues to mount specimens from confident and understand the basics, their his home in Pennsylvania. Last year the judgment is respected for many aspects Qing-Jun Li, Xishguangbanna Tropical volunteers mounted 20 percent of the of the mounting task. They are quite Botanical Garden, Chinese Academy of annual total, working, on average, 185 independent and know what needs to be Sciences; Alpinia (Zingiberaceae) hours per month. This has helped to fill done and how to do it. There is continued flowering (12/15-2/28). the shortage of staff to handle the oversight, but there is a trust between tremendous backlog and influx of volunteer mounter and supervisor that Andrew Henderson, New York Botanical specimens needing to be processed. constantly develops and grows, and the Garden; Arecaceae (12/27). Page 8 Profile Washington area in 1953 and was a of species concepts that were included. Continued from page 1 professor at George Washington Univer- Time and subsequent molecular work, sity. She worked in the Helenieae and did however, have largely supported Robin- at the Smithsonian was “inadequate;” many identifications especially in the son’s work. Robinson has done generic however, he spent a great deal of time years immediately following the death of level revisions of the Eupatorieae (with working with material from US and doing Blake. She was the advisor of two taxono- King), Liabeae, Heliantheae, and Vernoni- field work. A search on IPNI finds his mists that were eventually hired at US. eae. More recently he has helped imple- name associated with 4,159 plant names, Robert M. King (1929- ), a professor ment the presentation of a similar work of about 3,000 of which are in the Composi- at Catonsville Community College, came Guy Nesom in the Astereae. He has tae. He worked in the DC area from to the Washington area in 1962. During revised tribal limits and described new 1917-1959 and he added many specimens his time in the DC area he made many tribes and subtribes. The Liabeae had to the US collection. It is to Blake we field trips that resulted in numerous traditionally been recognized as one owe the wonderful collection of photos collections for US. He collaborated with unnatural core genus with three other of types from Europe that we all enjoy. Harold Robinson in the work on the genera, all in separate tribes, Senecion- He spent 42 years of his life producing Eupatorieae. King left the Washington eae, Heliantheae, Helenieae and Mutisi- 300 botanical papers, mostly on the area in 1997. One of King’s major eae. The Eupatorieae and Vernonieae, sunflowers and mostly in his spare time contributions to Compositae systematics traditionally placed together, were shown and he died at his desk on 31 December was introducing Harold Robinson to the to be only distantly related. It can safely 1959 (age 67). After his death, his family. be said that the work of H. Robinson has personal herbarium and library were Harold Robinson came to US in 1962 changed the way we do Compositae purchased by Lundell and ended up at the as a bryologist and moved over to working systematics. One particular series of University of Texas Plant Resource on the Compositae in 1964, applying results was the dismissal of the Cronquist Center in Austin. bryological techniques to a group that had view that the Rubiaceae were the closest Blake overlapped with José Cuatre- previously had inadequate anatomical relatives of the Compositae and that the casas who worked in the US herbarium study. Although he has rarely collected in Heliantheae were the primitive core until a few days before his death at the age the family, he has identified more than Continued on page 10 of 93. His career extended from his 30,000 specimens that he has received as studies in Barcelona and Madrid (1924- a “gift for determination” and is the 1931), graduate work in Berlin (1930-31) author or coauthor on over 600 papers where he knew Adolph Engler, Director- dealing with the family. His most recent ship of the Sección de Flora Tropical del accomplishments include the Heliantheae Jardin Botánico de Madrid (1933- 39), for the Flora of Ecuador. He is currently the years in Colombia (1939-47), in finishing the monograph on the Espeleti- Chicago (1947-1955) to his 41 years in inae by Cuatrecasas. Washington (1955-1996). Although The 1873 treatment by George Cuatrecasas (1903-1996) worked on Bentham divided the family into 13 tribes many families his concentration was on (the most frequently used suprageneric the Compositae of the northern Andes. He rank) which remained more or less the did a large amount of field work, mostly same until the 1980s. Many of the tribes in Colombia. He was an author or coau- had a large genus that provided the central thor of two subtribes and over 20 genera focus for the tribe: Senecio of the Senec- in five tribes. A quick check of IPNI ioneae, Eupatorium of the Eupatorieae, shows that 1,307 records from Cuatre- Aster of the Astereae, and Vernonia of casas are in the Compositae. At the time the Vernonieae. These genera remained of a 1985 festschrift, Cuatrecasas was largely unchanged having only a few listed with Robinson, Blake, Angel L. satellite genera removed from them. Cabrera, and Hermann Merxmüller, as Beginning with his work on the Eupatori- individuals who had made truly construc- eae, Robinson has tackled one large tive and insightful contributions to the genera after another. While his Eupatori- study of the Compositae in the first three eae work was done with King, subsequent quarters of the 20th century: three of the work has been done by himself or with top five synantherologists had worked at others. His detailed work resulted in the or with the collections from US. Cuatre- break up of several of these large genera. casas’ specimens, research materials and Each time the work was resisted because photographs remain at the Smithsonian others liked the convenience of a very Institution; his library is in Barcelona. few generic names that could be used for A. Head showing ray and disc florets, Kitty Parker (1910-1994), a student most specimens without concern for B. Disc floret showing pollen presen- of Lyman Benson in Arizona, came to the relationships or the identity of thousands tation, C. (cypsela) showing pappus. (Drawings by Alice Tangerini)

Page 9 Compositae published and unpublished trees for etic subfamilies are groups that used to be Continued from page 9 individual clades (usually tribes). This included in the now paraphyletic subfam- type of a supertree is a meta-tree, a “tree ily Cichorioideae (ca. 35 percent of the group, a concept referred to by Robinson of trees,” and it illustrates the current species in the family). The new classifica- as the “Heliocentric Theory of the thinking about the relationships among tions recognize 10 subfamilies and 35 Compositae.” the major tribes and subfamilies in the tribes, some new and some previously In 1980 the newest member of the Compositae. The basal group, which is described; it remains to be seen whether Compositae crew joined the Smithsonian. monophyletic and the sister-group to the or not this new classification will be Vicki A. Funk currently specializes in rest of the family, is the distinctive sub- accepted by the Compositae community. some of the tribes near the base or mid- family which contains The supertree makes it possible to level of the phylogeny of the family: less than 1 percent of the species in the look at the family as a whole and to try to Mutisieae, Liabeae, and Arctoteae. She family. Also monophyletic is the highly discern its origin and history as well as has an interest in phylogenetics and nested subfamily Asteroideae, which gain insights on character evolution (the biogeography and coordinates the super- contains ca. 65 percent of the species in figure below is a reduced version of the tree of the family. She does identifica- the family. Between the two monophyl- supertree). It is also an excellent method tions in her areas of interest and has collected extensively in Mexico, the Andes, and South Africa. Through the years a number of synantherologists have passed through the herbarium and the collections have benefited from their stay. Some have stayed long enough or visited often enough to annotate a number of speci- mens, including Walter Holmes, Rogers McVaugh, John Pruski, and Gisela Sancho. In addition, some curators at US, such as Conrad Morton and Warren L. Wagner, have shown an interest in the family even through it was not their area of specialization. olecular data came to Compos- ite systematics in the late M1980s and early 1990s and it was based on the work by Robert Jansen (TEX) and his collaborators. These individuals have literally turned the Compositae phylogeny upside down, showing that elements previously placed in the Mutisieae were a basal branch of the family and that the tribe Heliantheae s.l. was nested far up in the tree. Further- more, their work confirmed that the Vernonieae and Eupatorieae were actually in separate parts of the tree. Bruce Baldwin (UC) and his collaborators have proposed new suprageneric ranks for the Heliantheae s.l. and Jose Panero (TEX) and Funk have done the same for the base of the tree. Harold Robinson has pro- posed two supertribes for the Asteroid- eae. More recently Funk (along with 11 co- authors) has created a supertree for the Synantherologists at the US National Herbarium (clockwise from upper left): family that includes approximately 500 Sidney F. Blake (date unknown, photo from the Washington Biologists Field genera. The authors used a published tree Club Archives); José Cuatrecasas (date unknown, photo from the Botany Depart- for the backbone or base tree of the ment Archives); Kitty Parker (1973; photo from the NMNH Archives); Harold family and grafted onto the base tree the Robinson (2003; photo by M. Sangrey); Vicki Funk in Namaqualand, South Africa with Gazania (2002; photo by M. Koekemoer, PRE).

Page 10 Bawa, K.S., W.J. Kress, N.M. Nadkarni, S. Lele, P.H. Raven, D.H. Janzen, A.E. Lugo, P. Ashton, and T.E. Lovejoy. 2004. Tropical ecosystems into the 21st century. Science 306: 227-228. Clubbe, C., M. Gillman, P. Acevedo- Rodríguez, and R. Walker. 2004. Abun- dance, distribution and conservation significance of regionally endemic plant species on Anegada, British Virgin Islands. Oryx 38(3): 342-346. Dorr, L.J. and T. Wendt. 2004. A new species of Mortoniodendron (Malvaceae sens. lat.) from the rain forests of the Isthmus of Tehuantepec, Mexico. Lundellia 7: 44-52. Feuillet, C. 2004. Biodiversity in the Guianas and Venezuelan Guayana; data from the Boraginaceae, Gesneriaceae, and Passifloraceae. Flora of the Guianas Newsletter 14: 83-92. The reduced Compositae supertree showing tribes and a few other taxa. Feuillet, C. 2004. Passiflora phellos, a for determining critical areas of the tree growing in open areas), highly poisonous new species in subgenus Passiflora for future work. The creation of the (in its chemistry), fecund (not picky about (Passifloraceae). Novon 14(3): 285-287. supertree was an outgrowth of a new pollinators and have high seed set), and Funk, V.A., R. Chan, and S.C. Keeley. organization “The International Compos- agile (adapt to new edaphic conditions 2004 Insights into the evolution of the itae Alliance” (TICA) that had its first rapidly). As a result they are disgustingly tribe Arctoteae (Compositae: subfamily international meeting in Pretoria, January successful and have conquered the world. Cichorioideae s.s.) using trnL-F, ndhF, 2003, and the second meeting is to be The size and complexity of the family and ITS. Taxon 53(3): 637-655. held in June 2006, in Barcelona, Spain often result in taxonomists avoiding them. (hosted by Alfonso Susanna and Nuria They are often maligned by those who do Hawks, C., K. Makos, D. Bell, P.F. Garcia-Jacas). The organization is not study them; but maybe those who Wambach, and G.E. Burroughs. 2004. An supported by its website , hosted by Torsten vapor in herbarium cabinets. Taxon 53(3): Eriksson from Bergius Botanical Garden 783-790. (Stockholm) and uses The Compositae Newsletter (edited by Bertil Nordenstam, Kress, W.J. 2004. Paper floras: how long S) as a venue for publishing abstracts, and will they last? A review of Flowering other documents. The goal of the 2006 Plants of the Neotropics. American meeting is to produce a multi-volume Journal of Botany 91: 2124-2127. work on the family. Scientists from Kress, W.J., A.-Z. Liu, M. Newman, and around the world are expected to partici- Q.-J. Li. 2005. The molecular phylogeny pate and contribute to the publication. of Alpinia (Zingiberaceae): a complex TICA is run as an email society and and polyphyletic genus of gingers. anyone can join by sending a message to American Journal of Botany 92: 167- [email protected]. 178. We are often asked, “Why is the Compositae so large and successful while Price, J. P. and W. L. Wagner. 2004. its most closely related families, Speciation in Hawaiian angiosperm Calyceraceae and Goodeneacea, are small lineages: cause, consequence, and mode. with restricted distributions?” Well, it is Publications Evolution 58(10): 2185-2200. most likely the same old story: the Robinson, H. and A.J. Moore. 2004. New Comps are the new kid on the block. They Bawa, K.S., W.J. Kress, N. Nadkarni, and species and new combinations in are comparatively recently evolved (ca. S. Lele. 2004. Beyond paradise - meeting Rhysolepis (Heliantheae: Asteraceae). 50 million years ago), overly big (with the challenge in tropical biology in the Proceedings of the Biological Society of genera and species), opportunistic (often st 21 century. Biotropica 36: 437-446. Washington 117(3): 423-446.

Page 11 Art by Alice Tangerini

Stramentopappus pooleae (B. Turner) H. Robinson & V. Funk

The genus Stramentopappus was described by Robinson and Funk (Bot. Jahrb. Systematic 108: 213- 228. 1987) based on its lack of a sclerified pap- pus base and the reduc- tion of the outer ring of callus. The genus has the unusual chromosome number of n = 19 (most new world Vernonieae have chromosome counts of n = 17). This count was later found to be present in the two most closely related genera Leoboldia and Lepidonia.

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