Special Symposium Issue continues on page 14
Department of Botany & the U.S. National Herbarium The Plant Press
New Series - Vol. 19 - No. 3 July-September 2016 Botany Profile Green Things and Pollinator Wings: Symposium Explores Pollination Biology By Gary A. Krupnick ollination biology was the focus Professor of Biology from the University plant diversity. of the 14th Smithsonian Botanical of Massachusetts, was commended for his Droege said that while there are few PSymposium, held 20 May 2016 at extensive contributions to tropical biology, plant species that are pollinator special- the National Museum of Natural History international conservation, and pollina- ists, there are many bee species that are (NMNH) and the United States Botanic tion biology. In his acceptance speech pollen specialists. Specialist bee larvae Garden (USBG) in Washington, DC. Bawa expressed often fail Titled, “Bats, Bees, Birds, Butterflies his appreciation to thrive on and Bouquets: New Research in Pol- and gratitude to the pollen of lination Biology,” the meeting featured the Smithsonian other plants seven invited speakers, a poster session, Institution and and are often and an evening reception. The confer- the selection restricted to ence brought together over 340 biolo- committee. He was delighted to see so target plant blooming periods. Popula- gists, ecologists, government officials, many friends and colleagues and gave tion sizes tend to be low for special- horticulturalists, and master gardeners to thanks to all who have inspired him. ists which might lead to conservation celebrate the progress of pollination biol- Both Laurence Dorr and Seán Brady concerns. Droege’s mid-Atlantic bee ogy and to address issues of pollinator (Curator of Bees, NMNH) served as collection shows about 20 percent as loss and conservation. conveners. pollen specialists, with the most com- The Symposium began with opening mon being Andrena erigeniae (a Spring remarks by Maureen Kearney (Associate he first speaker was Sam Droege Beauty specialist), Ptilothrix bombi- Director for Science, NMNH) and Lau- (USGS Patuxent Wildlife Research formis (Hibiscus/Mallow specialist), and rence Dorr (Chair of Botany, NMNH). TCenter) who spoke about “Patterns Andrena violae (Violet specialist). The Kearney praised the partnership between in pollen and plant specialization among plant families with the most special- NMNH and USBG, and mentioned how native bees in eastern North America.” ists include Asteraceae, Ericaceae, and the symposium adds to the intellectual Droege’s macro photographs of bee Salicaceae. Specialist bees tend to be environment of the museum which often portraits from his online reference catalog the most vulnerable due to habitat loss, exercises an interdisciplinary research provided the backdrop for his talk about fragmentation, invasive species, and her- approach with a deep organismal focus in pollination from the bee’s point of view. bicide usage. Non-native plant species systematics, ecology, evolutionary biol- First he spoke about the high diversity of mostly favor generalist bee species, and ogy, and behavior. bee species in the United States – about thus Droege encourages the fostering of After the opening remarks, Ken- 4,000 species. He pointed out that there native habitat. neth Wurdack (Curator and Cuatrecasas are nearly as many bee species in Prince David Roubik (Smithsonian Tropical Committee Chair, NMNH) presented the George’s County, Maryland (249 species) Research Institute) next asked the ques- 14th José Cuatrecasas Medal in Tropical as there are in the United Kingdom (250 tion, “Whose bees are these? The pollen Botany to Kamaljit S. Bawa (see related species). He explained that 250 million taxonomy of bee nests and its story in story on page 14). This prestigious award years of evolution has led to complex recent decades.” Roubik explained that is presented annually to a scholar who and varied floral bee designs, nutritious pollen grains are nearly indestructible, has contributed significantly to advanc- pollen, complex secondary compounds, taxonomically informative, and open to and complex bee communities that reflect ing the field of tropical botany. Bawa, a Continued on page 17 Travel Vicki Funk traveled to Berlin, W. John Kress traveled to Costa Rica Jun Wen traveled throughout east- Germany (6/16 – 6/26) to be a plenary (6/9 – 6/16) to chair the Las Cruces Advi- ern China (6/23 – 7/22) to conduct field speaker at the Global Genome Biodiver- sory Committee meeting of the Organiza- research. sity Network conference, and to present tion for Tropical Studies. Kenneth Wurdack traveled to a workshop on Global Genome Initiative Gary Krupnick traveled to St. Louis, Wicomico County, Maryland (6/2) to (GGI) - Gardens (co-presented by Mor- Missouri (6/27 – 6/30) to present a talk collect Litsea aestivalis (Lauraceae) with gan Gostel). entitled “#EarthOptimism: Success Sto- Wesley Knapp (Maryland Department of Erika Gardner, Carol Kelloff, ries in Plant Conservation” and a poster Natural Resources) at Wetipquin Pond Melinda Peters, and Meghann Toner entitled “A Continental Approach to Preserve. traveled to Berlin, Germany (6/19 – 6/27) Orchid Conservation and Restoration” at Elizabeth Zimmer traveled to Austin, to attend the annual meeting of the Society the Global Partnership for Plant Conserva- Texas (6/17 – 6/22) to present a talk for the Preservation of Natural History tion conference at the Missouri Botanical entitled “DNA sequences identify cryptic Collections, where Gardner and Toner Garden. species of quillworts (Isoetes L.)” at the each presented a talk and Peters presented Sylvia Orli traveled to Amsterdam, Evolution 2016 meeting, and to attend a poster. Netherlands (4/30 – 5/5) with the Smithso- an editorial board meeting of Molecular Carol Kelloff and Mark Strong nian Digitization Program Office to meet Phylogenetics and Evolution as Deputy traveled to Zuni, Virginia (5/21 – 5/27) with Picturae and Naturalis to discuss the Editor-in-Chief. to collect rare plants in the Longleaf Pine specimen digitization project. communities for the Global Genome Ini- tiative (GGI) project. Nancy Khan traveled to Front Royal, Visitors Virginia (5/11 – 5/12) to attend a session of the Emerging Leaders Development Monica Carlsen, Missouri Botanical Gar- Yaling Wang, Xi’an Botanical Garden, Program at the Smithsonian Conservation den; Araceae and Zingiberales (2/17/15 China, and Shouzhou Zhang, Fairy Lake Biology Institute (SCBI). - 2/16/17). Botanical Garden, China; Global Genome Initiative Gardens Program partnership Gouri Mahadwar, University of Roch- The Plant Press meeting, Magnolia (Magnoliaceae) (4/12 ester; Legume phylogenetics internship - 4/14). New Series - Vol. 19 - No. 3 (9/1/15 - 8/11/16). Daniel Geiger, Santa Barbara Museum of Chair of Botany Morgan Gostel, George Mason Uni- Natural History; Oberonia (Orchidaceae) Laurence J. Dorr versity; Compositae and GGI-Gardens (4/15 - 4/19). ([email protected]) Program (9/1/15 - 8/31/17). Marcelo Devecchi, Universidade de São EDITORIAL STAFF George Wang, University of Louisiana at Paulo, Brazil; Simaba (Simaroubaceae) Lafayette; Plant-insect interactions (1/7 - (4/21 - 4/22) Editor 5/31). Gary Krupnick Lucas Jordao, Jardim Botanico de Rio de ([email protected]) Tomas Fer, Charles University, Czech Janeiro, Brazil; Mimosa (Fabaceae) (4/21 Republic; Zingiberales (2/1 - 6/3). Copy Editors – 4/22). Robin Everly, Bernadette Gibbons, and Joel Cohen, Rockville, Maryland; Plant Rose Gulledge Zvezdana Ukropina-Crawford, Berry DNA barcode internship (3/28 - 4/1). College; Crataegus (Rosaceae) (4/22). News Contacts Cassandra Quave, Emory University; Michele Conrad and 25 students, Mans- MaryAnn Apicelli, Rusty Russell, Alice Mediterranean ethnobotany (4/1). Tangerini, and Elizabeth Zimmer field University; Herbarium tour (4/23). Holly Summers, Cornell University; The Plant Press is a quarterly publication pro- Kate Wagle, University of Oregon; vided free of charge. To receive notification of Oenothera (Onagraceae) (4/5) when new pdf issues are posted to the web, please Botanical Art Collection (4/25 - 4/29). subscribe to the listserve by sending a message Chris Holder and Elizabeth Striano, to [email protected] containing only the Tom Lovejoy and 9 students, George following in the body of the text: SUBSCRIBE George Mason University, and Nancy Mason University; Plant conservation and PLANTPRESS-NEWS Firstname Lastname. Rybicki, U.S. Geological Survey; Trapa Replace “Firstname Lastname” with your name. herbarium tour (5/2). (Trapaceae) (4/6 - 4/7; 4/12 - 4/13). If you would like to be added to the hard-copy Marjorie Falanruw, U.S. Forest Service; mailing list, please contact Dr. Gary Krupnick at: Cintia Luz, Universidade de São Paulo, Department of Botany, Smithsonian Institution, Micronesian flora (5/3). PO Box 37012, NMNH MRC-166, Washington, Brazil; Schinus (Anacardiaceae) (4/11 - DC 20013-7012, or by E-mail: krupnickg@ 4/13). Kathleen Kron, Wake Forest University; si.edu. West Indian Ericaceae (5/11 - 5/12). Rowan Schley, Royal Botanic Gardens, Web site: http://botany.si.edu/ Kew; Brownea (Fabaceae) (4/11 - 4/14). Continued on page 10 Page 2 Taking Stock t has been an eventful year. Last summer when it average size of the 3,200 institutional herbaria in the Chair became apparent that I would likely become Chair world is a modest 100,000 specimens! As with the Iof the Department of Botany I realized that among symposium, this project also is a partnership. The many other things I would have to organize and orches- Digitization Program Office (DPO) of the Smithso- With trate a spring symposium. A number of ideas floated nian Institution has been a driving force, our princi- through my head and suppressing panic, the strongest pal benefactor, and the administrative contact with of all the ideas, I decided that a symposium on pol- Picturae, the Dutch company that owns and runs the lination biology might be interesting. I had dabbled conveyor belt and that subcontracts the transcrip- A with the subject early in my career and I knew that tions being done in Suriname. The very hard work in even though pollination biology per se is not a focus getting us to this point has been that of not only the of research done here in the Museum, a symposium on contractors employed by Picturae, but also Botany View this topic could easily underscore the valuable sys- and Museum staff and contractors who planned the tematic and phylogenetic research done by our Botany project, prepared the specimens for digitization, and Entomology Departments and inform some of our verified the transcriptions, and who are now import- research on the evolution of plants and insects. When ing the data into our collections database. I pitched the idea to Susan Pell of the U.S. Botanic The past year has also seen a transformation in L.J. Garden, our partner in this enterprise, she agreed and as the U.S. National Herbarium, which although far they say the rest is history. Of course, a good number from complete, augers well for the future. The new Dorr of Botany staff worked very hard to make this hap- cases designed to house our bulky bamboo collec- pen and I think we all should be very pleased with the tions (see The Plant Press 19(2): 8. 2016) have been result: an intellectually stimulating day with one of the installed and the specimens transferred from the old largest registrations yet for our symposium series. cases to the new. We are poised as well to order new Another inherited project, which was planned cases for the ferns and for an area currently used before but inaugurated after I became Chair, is our digi- as our specimen sorting center and these will be in tization conveyor project (see The Plant Press 19(2): place before the end of the year. The sorting center 6, 7. 2016). The scale of the accomplishments here in will be relocated and this will provide room for us to the last ten months is phenomenal. As of last week, expand the Asteraceae. The new cabinetry was pur- 566,824 herbarium specimens have been digitized, chased with monies awarded by the Smithsonian’s their label data have been or are now being transcribed, Collections Care and Preservation Fund (CCPF) for and the specimen images and information will soon be which we are extremely grateful. A separate CCPF fully searchable on our website. This is currently the grant is helping us to process a large backlog of largest herbarium digitization project in the Western grass specimens that will free up space and gener- Hemisphere and to put the scale of what has been ate a significant amount of exchange material. A accomplished into perspective, I need only add that the separate, moderately controversial change was the reintegration of the D.C. (District of Columbia) Her- barium into the main collection. Space and the need to focus staff resources on other matters drove this decision. In addition, we also came to the conclu- sion that the most efficient organization of the U.S. National Herbarium is to have the least number of separate collections possible. Having taken stock of a few of our major activities where do we go now? There will be another Smithsonian Botanical Symposium in May 2017 and we have initiated a discussion with the U.S. Botanic Garden about the theme. We are continuing the digitization of the herbarium and are beginning to think about how we can use very large data sets to address and answer questions with herbarium specimens that we could not imagine doing before. We are reorganizing the phanerogamic por- tion of the herbarium as a prelude to converting our filing scheme to a modern one that better reflects our understanding of phylogeny. Finally, you might ask why these things matter. They matter because we are try- ing to support and improve our research on the systematics and evolu- tion of plants, fungi (i.e., lichens), and algae. Specimens from the United States National Herbarium are placed on a conveyer belt for rapid digitization. (photo by Ingrid Lin)
Page 3 Schuettpelz. Dal Forno received her Ph.D. the differentiation and diversification of Staff in Environmental Science and Public plant groups in some of the more chal- Policy from George Mason University. lenging environments of North America. Research & Her dissertation research focused on the Edwards has a Ph.D. in phylogenetics Activities systematics of tropical basidiolichens in and biogeography from The University of the Dictyonema sensu lato clade (Agari- Queensland in Australia studying dif- cales: Hygrophoraceae). Dal Forno was ferentiation, niche evolution, and species As a member of the Botanical Art Society awarded an NSF fellowship for research identity in a recent and complex radiation of the National Capital Region (BASNCR) using biological collections and will be of monsoon tropical trees, broadleaved Alice Tangerini has artwork on display in obtaining microbiome data from fresh and paperbarks (Melaleuca, Myrtaceae). He two of their recent exhibits in the Wash- historical lichen collections deposited at is also recovering from two years as a ington DC area. In the exhibit, “Botanica the U.S. National Herbarium. She will be Drosophila speciation genetics postdoc at 2016: The Art & Science of Plants,” at comparing these data in both space (from the University of Rochester (identifying Brookside Gardens in Wheaton, Maryland different localities) and time (at various the genetic mechanisms of hybrid incom- (June 16 - July15), Tangerini has exhib- ages). Her main objective is to test the patibility and the large X effect between ited “Tree Peony” in pen and ink made hypothesis that microbiome data from D. simulans and D. mauritiana), and is specifically for an adult coloring book of specimens deposited in herbaria represent enjoying supra-freezing temperatures, blue garden plants. At the opening reception, the original bacterial assemblage associ- skies and dabbling with chlorophyll again. Tangerini presented a lecture on botani- ated with the living lichen thallus. This His research can be followed at http:// cal illustration techniques. In “Off the opens up a remarkable opportunity to add bortedwards.weebly.com/ and on Twitter Beaten Path,” which features botanical another data layer to millions of existing and Instagram @BortEdwards. works in non-traditional presentation and collections deposited in museums world- composition at the Strathmore Mansion in wide. Laura Tancredi is the newest staff Bethesda, Maryland (June 11 - July 31), member of the Department of Botany. Tangerini has exhibited a mixed media Bort (Robert) Edwards arrived in As part of the Information Management piece, “Tulips.” January as a Powell Center Postdoctoral team, her main roles will be to enhance the Fellow under the guidance of Vicki Funk. Botany specimen catalog through digitiz- The Powell Center is a USGS initiative ing, inventorying and imaging specimens, for collaborative analysis and synthesis performing quality control for the rapid and Edwards’ project brings together digitization project, and supervising data and investigators from SI, USGS, volunteers and interns who will assist with USDA, CSIRO, NSF, and a number of the improvement of the online records. other organizations and universities. They Before coming to the Smithsonian as a New Faces aim to leverage big data in the form of Collections Program Technician (CPT) geochemistry and geology, climate, plant in 2012, Tancredi worked as an archae- Manuela Dal Forno joined the Botany distributions, and sequence data to inves- ologist at George Washington’s Mount Department as a postdoctoral fel- tigate whether the evolution of extreme Vernon. She therefore comes to Botany low in June 2016, working with Eric environmental tolerances has allowed with plenty of experience “collecting” in
New faces in the Department of Botany (from left): Manuela Dal Forno, Bort Edwards, Laura Tancredi. Page 4 the dirt. Tancredi’s first project as a CPT, and first exposure to the world of pressed specimens, was with the National Anthro- pological Archives, mounting ethnobotani- cal specimens as part of the Recovering Voices Initiative. She subsequently spent over a year in the Division of Fishes cata- loging ~15,000 specimen lots in their fluid collection, and moving and rehousing the osteological collection. She participated in the Department of Entomology’s species level inventory, catalogued the Department of Paleobiology’s Amber Collection, and spent about eight months last spring and summer with the Department of Botany preparing specimens for the conveyor belt digitization project. Awards & Grants
The Journal of Systematic Evolution (JSE) recognized the 2015 paper, “A worldwide phylogenetic classification of the Poaceae (Gramineae),” by Robert Soreng, Paul Peterson, Kostyantyn Romashchenko, Gerrit Davidse, Fernando O. Zuloaga, Emmet J. Judziewicz, Tarciso S. Filguei- ras, Jerrold I. Davis, and Osvaldo Mor- Volunteers from the Department of Botany: (1) Ardith and Jim Harle both rone as one of four JSE Outstanding volunteer in Botany. Jim manages the map project, which has expanded across the Papers (2014 and 2015). The 2015 paper, museum, and Ardith works on the plant mounting opportunity. (2) The Monday “Expression patterns of AP1, FUL, FT and plant mounting crew (clockwise from top left): Kat Rankin (retired plant mounting LEAFY orthologs in Vitaceae support the coordinator), Irmgard Dugge, Carol Pihstrom, Gwen Petitjean, Jo Ann McCann, homology of tendrils and inflorescences and Bayla White. (3) Larry and Alice Owens with Melinda Peters. Larry works on throughout the grape family,” by Ning the plant mounting opportunity. (4) Mary Monsma in the studio scanning her fern Zhang, Jun Wen, and Elizabeth Zim- illustrations. mer, was recognized as one of four JSE Outstanding Papers by Young Investiga- The museum held a special event in published, as the classification of ferns tors (2014 and 2015). Both papers were April to honor the volunteers and to give had changed too much. In 2012 Monsma nominated by JSE editors and were our thanks. During the event a number of returned to Botany to begin a long term selected largely based on the citations per Botany volunteers were recognized for volunteer project to scan, inventory year. In addition to a certificate, winners their years of service: 5 years (Ann Car- and organize her fern illustrations from received a cash award. doni, Deanna Crouse, Kathleen Griffin, her contract years. With her additional Ardith Harle), 10 years (Cynthia Ford, knowledge in Photoshop she has been able Rosalie Elliott, Bayla White), 20 years to reconstruct, label and optimize the illus- Volunteer (Larry Owens), 25 years (Margaret Sch- trations for future inclusion in Botany’s Appreciation in weitzer), and 30 years (Gwen Petitjean). public website. Mary Monsma was selected from the Botany Department of Botany for the “Above April was designated as Volunteer and Beyond 2016” award. Monsma first Appreciation Month at the National worked in Botany in the early 1970s as Museum of Natural History. The Depart- a contract illustrator drawing ferns for ment of Botany has many dedicated David Lellinger and his publication, volunteers involved in all aspects of our The Ferns and Fern-allies of Costa Rica, work. From plant preparation to map Panama, and the Choco: Part 1. In 30 curating and data entry, this extraordinary years she completed 582 drawings for that and talented group of volunteers deserves publication, and another 596 drawings special recognition. for the second volume. Part 2 was never
Page 5 Dan Henry Nicolson (1933-2016) Dan Nicolson was born on 5 Septem- ber 1933 in Kansas City, Missouri and grew up in Shenandoah, Iowa where his parents ran the Henry Field’s Seed and Nursery Company, which was then one of the largest mail order seed companies in the country. After graduating from Shenandoah High School (1951) Nicolson enrolled in Grinnell College, Grinnell, Iowa. Four years later, not only did he graduate with a B.A. (1955) but he also published his first botanical paper: a revi- sion of the genus Asclepias (milkweed) in Iowa. Evidently there was an expectation that he would take over the family busi- ness and he dutifully pursued a M.B.A. (1957) at Stanford. Botany, however, proved to be more alluring than business and rather than return to Shenandoah, he moved further east to pursue gradu- ate study in Botany at Cornell University (M.S., 1959 and Ph.D., 1964). While studying at Cornell, Nicolson met and married Alice (Allie) Black Craw- ford. The newly wedded couple attended the 1959 International Botanical Congress in Montreal for their honeymoon. This unorthodox destination must not have bothered his wife too much because they remained married for 57 years. While in graduate school Dan and Allie spent almost two years (1960-62) in Malaysia and Southeast Asia where he did the field work that led to a monograph of the genus Aglaonema (Araceae), the subject of his dissertation. In 1964 Nicolson was hired by the Smithsonian Institution to fill a vacancy created when Richard (Dick) Cowan was Nicolson continued his focus on involved in botanical nomenclature and promoted to Assistant Director of the U.S. Asian botany and spent a year (1966-67) in the International Association for Plant National Museum (now National Museum collecting plants in Nepal as a Senior Taxonomy (IAPT) so much so that for of Natural History). During his first year Fulbright Fellow. This was followed by close to a generation Dan Nicolson and on the job Nicolson attended the Interna- work in India (1968-74) where he spent Botanical nomenclature were almost tional Botanical Congress in Edinburgh three months of each year in Bangalore synonymous. He served (1979-99) as and spent three months collecting on the (now Bengalaru) collaborating with Father nomenclature editor of the journal Taxon island of Dominica in the Caribbean. The C.J. Saldanha on the flora of the Hassan and was (1981-2006) also a member of the following year as the junior curator in the District, which was published in 1975. He editorial committee for the International department, he was tasked with organizing also spent a month in Sri Lanka in 1979 Code of Botanical Nomenclature helping the move of the U.S. National Herbarium and later contributed a treatment of the to prepare five separate editions. He held from the Castle to its current location Araceae to the multi-volume A revised numerous offices in the IAPT including in the West Wing of the Natural History handbook of the flora of Ceylon, which president (1993-99). In conjunction with Building. Several years later he organized was then being published for the Smithso- his interest in nomenclature, Nicolson also the Herbarium Services Unit (now Core nian Institution and the National Science delved deeply into Botanical bibliography Collections Management) and was also Foundation. A three-month long trip to and history. His most important contribu- instrumental in recruiting Ruth Schallert to Yunnan, China in 1983 was Nicolson’s last tions to these last fields wereAn interpre- serve as the first librarian for what is now extensive foray into the field. tation of Van Rheede’s Hortus Malabari- the Botany and Horticulture Library. In the 1970s Nicolson became deeply cus (1988), The Forsters and the botany of
Page 6 the second Cook Expedition (1772-1775) (2004), and the final two supplements to Taxonomic Literature, ed. 2 (2008, 2009). All four of these books were published with an IAPT imprint. Nicolson liked to quip that his sci- entific work could be divided into three areas: 1, Monographic work in Araceae (knowing a lot about little); 2, Floristic work in India and the West Indies (know- ing a little about a lot); and 3, Nomen- clature (knowing a little about really nothing). He was perhaps a little too self- effacing. His contributions to Botany were important and he received recognition for them: a Science Achievement Award (2004) from NMNH, the Stafleu Medal (2005) from the IAPT, and the Secretary’s Research Award (2009) from the Smithso- nian Institution. In 2005 Nicolson attended the Inter- national Botanical Congress in Vienna, his ninth and final congress. Heretire d in December of the same year and initially devoted his time to bringing the monu- mental IAPT-sponsored project Taxonomic Literature to a conclusion. Health issues, however, prevented him from taking on additional projects. In addition to his wife Allie, Nicolson’s immediate survivors are his three children, John, Sally, and David. David works in the Natural History Building as a Data Devel- opment Coordinator for the Integrated Taxonomic Information System (ITIS), which is part of the USGS. The Department of Botany plans to hold a Memorial Service for Nicolson in the early Fall.
Opposite page: Dan Nicolson, recipient of the Distinguished Career in Science Award given by the Washington Academy of Sciences, in 2008. This page, top to bottom: Nicolson supervises the move of the Department of Botany and the assembled collections to the new west wing of the Museum of Natural History, in 1965. Nicolson displays a specimen during the blooming of Amorphophallus titanium at the United States Botanic Garden, in 2003. Nicolson receives the Stafleu Medal from Pieter Baas during the 17th International Botanical Congress in Vienna, Austria, in 2005. Page 7 Smithsonian Scientists Participate in National BioBlitz In celebration of the Centennial of the National Park Service (NPS), the Smithsonian Institution joined NPS and the National Geographic Society on May 20-21 to participate in the 2016 National BioBlitz. The event included all-taxa rapid biological surveys, in which teams of vol- unteers, each led by subject matter experts, found and identified as many species as possible in national parks throughout the country. A concurrent Biodiversity Festi- val with hands-on learning activities took place on the National Mall at Constitution Gardens. Amorpha fruticosa (left) and Persicaria arifolia (right) were both observed at Dyke This two-day event centered around Marsh Wildlife Preserve in Virginia during the 2016 National BioBlitz. (photo by a 24-hour BioBlitz inventory period Nancy Khan) in which teams of scientists, teachers, children, and other community members Over nine thousand species in differ- Washington area. Representing Smithso- worked together on a series of different ent taxonomic groups, and over 85,000 nian’s Department of Botany was Nancy inventories to discover and document as observations were documented, and tens Khan who led a plant survey of Dyke many species of plants, animals, microbes, of thousands of public participants were Marsh Wildlife Preserve, and Manuela fungi, and other organisms as possible. engaged in our national parks. Among the Dal Forno who led a lichen inventory on The primary data collection tool for all taxonomic groups, 40,378 plant observa- Theodore Roosevelt Island. BioBlitzes was iNaturalist
Sharon and Michael Rosenthal show the type specimen of the quillwort Isoëtes viridimontana. (photo Ken Wurdack) Page 10 Celebrating tors. Krupnick was one of the scientific advisors on the poster design. Pollinators and During the program, Krupnick also provided museum visitors information on Plants how to join the Million Pollinator Garden National Pollinator Week 2016 was Challenge, a way to register public and celebrated June 20-26 in an effort to raise private gardens and landscapes to support awareness of the decline in important pol- pollinators. The Million Pollinator Garden linator populations. The National Museum Challenge was launched by The National of Natural History held a number of events Pollinator Garden Network, an unprece- throughout the week, including “Expert is dented collaboration of national, regional, In!” programs throughout the museum and conservation and gardening groups, a Pollination Party in the Pollinator Gar- including Smithsonian Gardens and the den. Many of the events moved beyond National Museum of Natural History, to simply raising awareness, giving partici- support the President’s Executive Strategy pants opportunities to contribute towards to “Promote the Health of Honey Bees and pollinator protection. Other Pollinators.” On June 20, Smithsonian experts came On June 21, Smithsonian Gardens out to the museum galleries to talk about hosted its third annual Pollination Party. unique pollinators. Visitors of all ages The event was kicked off by the unveiling swarmed their tables to learn about the of the newly renamed Pollinator Garden behavior of plant pollinators. Gary Krup- outside the National Museum of Natural Gary Krupnick displays plant and nick explained to visitors the importance History. The original Butterfly Habitat bee specimens and talks about the of “Trees for Bees,” a topic highlighted in Garden was renamed to reflect the grow- importance of trees for native bees the 2016 Pollinator Week poster issued by ing importance of supporting pollinator during an “Expert is In” session at the the Pollinator Partnership. With their pro- health championed by the formation of National Museum of Natural History. fusion of flowers, trees are a convenient a task force by President Obama in 2014 volunteers to help make biodiversity data food source for bees and other pollinators. and the implementation of the Million more accessible by transcribing label Krupnick displayed specimens of tulip tree Pollinator Garden Challenge. Smithsonian information from museum specimens. (Liriodendron tulipifera), staghorn sumac Gardens’ reinterpretation of this space Krupnick, who represented the Smith- (Rhus typhina), flowering dogwood (Cor- educates visitors on the wide diversity sonian Institution on the Pollinator Health nus florida), and eastern redbud (Cercis of pollinators and the types of plants that Task Force, worked hard to ensure that canadensis), and shared stories of their support them. The garden display also the Smithsonian is well-represented in the natural history and importance to pollina- shows what can be done to create beautiful pollinator-friendly gardens. Strategy, which draws attention to the pol- On June 22, the White House released linator research, education, and landscapes the Pollinator Partnership Action Plan at the National Museum of Natural His- (PPAP), building on Federal actions to tory, Smithsonian Conservation Biology improve pollinator health by facilitat- Institute, National Zoological Park, Smith- ing additional state and private-sector sonian Environmental Research Center, engagement. The PPAP furthers President Smithsonian Tropical Research Institute, Obama’s June, 2014, memorandum that Smithsonian’s Forest Global Earth Obser- focused the attention of Federal agencies vatories, and Smithsonian Gardens. on the plight of the pollinators—honey bee colony mortality rates that impact the Rising CO2 Depletes viability of commercial beekeepers and Pollen’s Nutritional agricultural pollination services; monarch butterfly declines that threaten its iconic Potency, Bees Suffer continental-scale migration; and other pollinator species quietly slipping toward -Adapted from Smithsonian Insider extinction. The PPAP highlights ways in Unless you’ve been living under a rock which state, local, and tribal governments; for the last few years, you’ve doubtless the private, academic, and non-profit sec- caught at least a passing reference to the tors; and the general public can contribute plight of the beleaguered bee. Bees of all to achieving the goals of the national types pollinate an estimated 75 percent Strategy to Promote the Health of Honey of our fruit, nut and vegetable crops, and Bees and Other Pollinators. Among the they’ve been suffering population declines The 2016 Pollinator Week poster, “Trees highlights in the PPAP is how the Smith- in recent years from a variety of suspected for Bees.” sonian Institution is working to recruit Continued on page 12 Page 11 Pollen Continued from page 11 sources. These die-offs are a major con- cern to farmers and hobby beekeepers, as well as anyone who likes to eat. Bees like food too, but a new study led by the U.S. Department of Agriculture’s Agricultural Research Service has found that recent increases in carbon dioxide emissions over the last several decades have made a key autumnal food source less nutritious than in the past. This adds one more potential factor to the cocktail of stressors making bees vulnerable to attack by pests, pathogens and pesticides. Three of the many historical goldenrod records kept in the U.S. National USDA Agricultural Research Service Herbarium. These plants were collected in Arizona, Florida and Alabama (left plant physiologist Lewis Ziska, together to right). Goldenrod samples in this herbarium were used to measure how rising with colleagues from Purdue University, atmospheric CO2 has impacted protein in pollen. Williams College and the Smithsonian, compared the protein content of pol- protein content has declined by about one “The Smithsonian herbarium goes right len from historic specimens of Canada third since 1960, from 18 to 12 percent. to the heart of America’s past, because goldenrod (Solidago canadensis) from the A field experiment that simulated when it started our first collections were a similar range of CO exposures, but from people like Lewis and Clark, fron- U.S. National Herbarium collection at the 2 included future CO concentrations up to tiersmen who were identifying what was Smithsonian’s National Museum of Natu- 2 ral History with pollen from field trials 500 parts per million (ppm), confirmed the unique to America. No European had seen simulating varying levels of atmospheric herbarium results. Still, as Ziska pointed what was there,” Clark said. “It’s a very out, it’s not clear if there’s an upper limit clear snapshot of what our natural environ- carbon dioxide. They found that as CO2 levels increased, the protein content of the to the effect, or if the protein content will ment looked like at that time.” eventually stabilize regardless of the CO Whether or not the bees will be able to pollen decreased. The study was published 2 in the journal Proceedings of the Royal concentration in the atmosphere. adjust to a degraded pollen source is still Society B (283: 20160414; 2016). The silver lining to the study, he said, unclear. Goldenrod, which blooms from July is that commercial and hobby honeybee “As with most biological organisms, through October across most of North keepers may be able to counteract any there is some capacity for adaptation, but America, is one of the most widely nutritional deficiencies by supplying how much is an open question,” Ziska available sources of pollen for late-year additional protein late in the fall. How to paused, then continued. “I often read ameliorate the impact on wild, solitary where ‘CO is plant food’, but that simple foraging. Though nectar is the bees’ main 2 food source during the warmer months, bees is, however, unclear. meme falls short of understanding the “We wanted to conduct a robust study complexity of CO in plant biology. I bees need the fats, vitamins and minerals 2 from pollen protein to make it through by looking at both the historical record suspect that plant pollinators are just the the winter. Because they only store small and field experiments,” Ziska says. tip of the iceberg—or flower.” amounts of it, fluctuations in the amount Though he started out by visiting her- or quality of the pollen itself can directly bariums around the world, including in Publications affect bee health. Australia, the Smithsonian’s herbarium “Goldenrod was the focus because it’s “has by far the best and most diverse set of Appelhans, M.S., S. Krohm, S. Manafza- the last source of pollen for bees before herbarium samples for studying long term deh and J. Wen. 2016. Phylogenetic they overwinter,” Ziska says. “Whatever climate effects,” he adds. placement of Psilopeganum, a rare mono- happens to goldenrod in the fall can be a Andrew Clark, who now works for typic genus of Rutaceae (the citrus family) harbinger of overall bee health and their the USDA but at the time of the study endemic to China. J. Syst. Evol. http:// ability to survive until spring.” was a Smithsonian herbarium collec- dx.doi.org/10.1111/jse.12208 How rising CO affects insects’ food tions specialist guiding Ziska through the 2 Carlucci, M.B., G.D.S. Seger, D. Sheil, sources has never been investigated in Solidago specimens, says that the use of goldenrod in this way is a fairly novel I.L. Amaral, G.B. Chuyong, L.V. Ferreira, much depth, despite more than 100 such U. Galatti, J. Hurtado, D. Kenfack, D.C. studies on human food sources suggest- approach to the kinds of research typically conducted with herbariums. Pollen studies, Leal, S.L. Lewis, J.C. Lovett, A.R. Mar- ing the same kind of impact, Ziska said. shall, E. Martin, B. Mugerwa, P. Munishi, Using Smithsonian herbarium samples for example, aren’t done very often any more, and typically focus on the structure Á.C.A. Oliveira, J.C. Razafimahaimodi- dating from 1842 through to 1998, as well son, F. Rovero, M.N. Sainge, D. Thomas, as more recently collected samples from of the pollen itself to aid in plant family classification. V.D. Pillar and L.D.S. Duarte. 2016. other sources, Ziska found that pollen Phylogenetic composition and structure of
Page 12 tree communities shed light on historical PloS One 11(4): e0153565. http://dx.doi. the population dynamics of a threatened processes influencing tropical rainfor- org/10.1371/journal.pone.0153565 Caribbean dry forest cactus. Biol. Conserv. est diversity. Ecography http://dx.doi. 196: 156-164. http://dx.doi.org/10.1016/j. org/10.1111/ecog.02104 Katinas, L., M.P. Hernández, A.M. Aram- biocon.2016.02.015 barri and V.A. Funk. 2016. The origin of Costion, C.M., A.J. Lowe, M. Rossetto, the bifurcating style in Asteraceae (Com- Seberg, O., G. Droege, K. Barker, J.A. R.M. Kooyman, M.F. Breed, A. Ford and positae). Ann. Bot. 117(6): 1009-1021. Coddington, V.A. Funk, M. Gostel, G. D.M Crayn. 2016. Building a plant DNA http://dx.doi.org/10.1093/aob/mcw033 Petersen and P.P. Smith. 2016. Global barcode reference library for a diverse Genome Biodiversity Network: saving a tropical flora: an example from Queen- Li, R. and J. Wen. 2016. Phylogeny blueprint of the Tree of Life – a botani- sland, Australia. Diversity 8(1): 5. http:// and diversification of Chinese Arali- cal perspective. Ann. Bot. http://dx.doi. dx.doi.org/10.3390/d8010005 aceae based on nuclear and plastid DNA org/10.1093/aob/mcw121 sequence data. J. Syst. Evol. 54(4): 453- Culley, T.M., C.D. Huebner and A. Novy. 467. http://dx.doi.org/10.1111/jse.12196 Shi, W., J. Wen and B. Pan. 2016. A 2016. Regional and local genetic varia- comparison of ITS sequence data and tion in Japanese stiltgrass (Microstegium Martin, M.D., M.T. Olsen, J.A. morphology for Calligonum pumilum and vimineum). Invasive Plant Sci. Manag. Samaniego, E.A. Zimmer and M.T. Gil- C. mongolicum (Polygonaceae) and its 9(2):96-111. http://dx.doi.org/10.1614/ bert. 2016. The population genomic basis taxonomic implications. Phytotaxa 261(2): IPSM-D-15-00055.1 of geographic differentiation in North 157-167. http://dx.doi.org/10.11646/ American common ragweed (Ambrosia phytotaxa.261.2.5 Fernández-Alonso, J.L., P.A. Fryxell and artemisiifolia L.). Ecol. Evol. 6(11): 3760- L.J. Dorr. 2016. Malvaceae, pp. 1535- 3771. http://dx.doi.org/10.1002/ece3.2143 Strong, M.T. 2016. Three new Cyper- 1570. In R. Bernal, S.R. Gradstein and aceae from the Cuyuni-Mazaruni M. Celis (eds.). Catálogo de Plantas y McManus, K.M., G.P. Asner, R.E. Martin, Region of Guyana (South America). Líquenes de Colombia 2. Instituto de K.G. Dexter, W.J. Kress and C.B. Field. Novon 24(4): 401-407. http://dx.doi. Ciencias Naturales, Universidad Nacional 2016. Phylogenetic structure of foliar org/10.3417/2015032 de Colombia, Bogota. spectral traits in tropical forest canopies. Remote Sens. 8(3): 196. http://dx.doi. Thornhill, A.H., B.D. Mishler, N.J. Knerr, Feuillet, C. 2016. Two new combina- org/10.3390/rs8030196 C.E. González-Orozco, C.M. Costion, tions in Euploca Nutt. (Heliotropiaceae, D.M. Crayn, S.W. Laffan and J.T. Miller. Boraginales) and a conspectus of the spe- Ochyra, R. and R.R. Ireland. 2016. 2016. Continental-scale spatial phyloge- cies of the Guiana Shield area. PhytoKeys Isopterygium tenerifolium (Hypnaceae, netics of Australian angiosperms provides 61: 101-124. http://dx.doi.org/10.3897/ Bryophyta) - one more Afro-American insights into ecology, evolution and phytokeys.61.6260 disjunct. Herzogia 29(1): 72-78. conservation. J. Biogeogr. http://dx.doi. org/10.1111/jbi.12797 Feuillet, C. 2016. Lectotypifications Osuri, A.M., J. Ratnam, V. Varma, P. of six taxa in the Boraginales (Cor- Alvarez-Loayza, J. Hurtado Astaiza, M. Zhang, K.M., Y. Shen, X.Q. Zhou, Y.M. diaceae and Heliotropiaceae). PhytoKeys Bradford, C. Fletcher, M. Ndoundou- Fang, Y. Liu and L.Q. Ma. 2016. Photo- 62: 95-100. http://dx.doi.org/10.3897/ Hockemba, P.A. Jansen, D. Kenfack, A.R. synthetic electron-transfer reactions in the phytokeys.62.6259 Marshall, B.R. Ramesh, F. Rovero and gametophyte of Pteris multifida reveal M. Sankaran. 2016. Contrasting effects of the presence of allelopathic interference Harris, A.J., C. Fu, Q.Y. Xiang, L. Hol- defaunation on aboveground carbon stor- from the invasive plant species Bidens land and J. Wen. 2016. Testing the mono- age across the global tropics. Nature Com- pilosa. J. Photochem. Photobiol. B: phyly of Aesculus L. and Billia Peyr., mun. 7: 11351. http://dx.doi.org/10.1038/ 158: 81-88. http://dx.doi.org/10.1016/j. woody genera of tribe Hippocastaneae of ncomms11351 jphotobiol.2016.02.026 the Sapindaceae. Mol. Phylogenet. Evol. 102: 145-151. http://dx.doi.org/10.1016/j. Peterson, P.M. and R. Soreng. 2016. A Zhang, N., J. Wen and E.A. Zimmer. ympev.2016.06.001 revision of Poa subsection Aphanelytrum 2016. Another look at the phylogenetic (Poaceae, Pooideae, Poaeae, Poinae); and position of the grape order Vitales: Chlo- Henkel, T.W., C.L. Kelloff, S.N. Alexan- a new species, Poa auriculata. PhytoKeys roplast phylogenomics with an expanded der and V.A. Funk. 2016. Smithsonian 63: 107-125. http://dx.doi.org/10.3897/ sampling of key lineages. Mol. Phylo- plant collections, Guyana: 1992–2014, phytokeys.63.8198 genet. Evol. 101: 216-223. http://dx.doi. Terry W. Henkel. Smiths. Contrib. Bot. org/10.1016/j.ympev.2016.04.034 104: vi-206. http://dx.doi.org/10.5479/ Rabeler, R.K. and W.L. Wagner. si.1938-2812.104 2016. New combinations in Odon- Ziska, L.H., J.S. Pettis, J. Edwards, J.E. tostemma (Caryophyllaceae). PhytoKeys Hancock, M.B. Tomecek, A. Clark, J.S. Howard, M.G., W.J.F. McDonald, P.I. 63: 77-97. http://dx.doi.org/10.3897/ Dukes, I. Loladze and H.W. Polley. 2016. Forster, W.J. Kress, D. Erickson, D.P. phytokeys.63.8181 Rising atmospheric CO is reducing the Faith and A. Shapcott. 2016. Patterns 2 Rojas-Sandoval, J., E.J. Meléndez- protein concentration of a floral pollen of phylogenetic diversity of subtropical source essential for North American bees. rainforest of the Great Sandy Region, Aus- Ackerman and D. Anglés-Alcázar. 2016. Assessing the impact of grass invasion on Proc. R. Soc. B 283(1828): 20160414. tralia indicate long term climatic refugia. http://dx.doi.org/10.1098/rspb.2016.0414 Page 13 Bawa Receives 14th Cuatrecasas Medal The Department of Botany and the in consultation with other plant scientists contributions have addressed the chal- United States National Herbarium present outside of the Smithsonian Institution. lenges of biodiversity conservation and this award to a botanist and scholar of Nominations for the Medal are accepted sustainability, especially for India and international stature who has contributed from all scientists in the Botany Depart- its centers of biodiversity in the Western significantly to advancing the field of ment. The award consists of a bronze Ghats and Himalayas. In 1996 he founded tropical botany. The José Cuatrecasas medal bearing an image of José Cuatreca- the Ashoka Trust for Research in Ecology Medal for Excellence in Tropical Botany sas on the front with the recipient’s name and the Environment (ATREE), which is named in honor of Dr. José Cuatrecasas, and date of presentation on the back. has developed research and education a pioneering botanist and taxonomist, who Highlights from past presentations to the programs to address key environmen- spent nearly a half-century working in the recipients are available at
Sam Droege USGS Patuxent Wildlife Research Center Speakers, conveners, and award recipients of the 2016 Smithsonian Botanical “Patterns in pollen and plant special- Symposium at the National Museum of Natural History (from left): Ken Wurdack, ization among native bees in eastern North Laurence Dorr, Kamal Bawa, Robert Raguso, Tatyana Livshultz, David Roubik, America” Seán Brady, Candace Galen, Matthew Koski, and Nathan Muchhala. (photo by Tomas Fer) The long co-evolution of flowering plants and their insect pollinators has cre- brief mention of conservation implications dinal gradient on Pennsylvania Mountain ated an abundance of strategies on both for both plants and bees, and presentation (CO), past bumble bee host species flower sides of the plant/insect equation. Many of a number of examples of specialized for only 33% of the time span observed insects are visitors to flowers and their role bee/plant pairings. in the 1970s. Advancement of flowering as floral visitor from the plant perspective phenology with warming is especially varies from outright thievery to obligate Candace Galen pronounced in historically late season pollinator. Of those insect visitors, native University of Missouri host plants, leading to increased flowering overlap among species, causing temporal bees are almost entirely unique in that they “Shifting baselines and changing mismatches between bumble bee colony purposefully gather pollen and nectar to partners: Ecological and evolutionary production and resource availability, and provision nests for their young. The 800 responses to climate change in alpine favoring generalized foraging in two long- species of native bees east of the Missis- bumble bees and their host plants” sippi River in North America have greatly term resident bumble bee species. Results Bumble bees (Bombus) are a key polli- compartmentalized their plant interactions. from pollination experiments suggest that nator guild in North America with multiple About 20 to 25 percent are highly special- a positive feedback loop links general- species in decline. Globally, bumble bees ized, using pollen from at times only one ized foraging in bumble bees to foreign are moving upward in altitude, suggest- species or genus of plant. Other species pollen contamination and impaired seed ing alpine regions represent potential are less restrictive in the pollen they use, set of alpine host plants. Using herbarium thermal refugia from climate change in but still may largely focus on gathering specimens to reconstruct past trajectories lower, southern regions of their range. pollen from a single plant family and have of pollination quality we can test this idea, Yet our research over four decades in the only a brief, six-eight week period of flight addressing broad scale impacts of global Rocky Mountains indicates that as warm- activity. A smaller number of species, but warming on pollination services. ing continues, these high altitude refugia often the most dominant in terms of indi- are themselves likely to be imperiled. In viduals present, visit a wide range of plant Matthew Koski central Colorado (USA), encroachment families, often are social or colonial in life University of Virginia, USA of woody vegetation near timberline history strategy (but never to the extent (tundra greening) and reduced flower- “The evolutionary ecology of ultra- that the introduced honeybee has taken ing of tundra plant communities near the violet floral pigmentation at micro- and social living) and are active throughout the krummholz ecotone has led to landscape macroevolutionary scales” flowering season. Specialized structures scale declines in historically preferred The tremendous diversity in flower in both plants and their bee counterparts resources for bumble bee foragers. Only color has fascinated biologists for decades. are abundant and will be illustrated in this in small habitat patches near alpine sum- Because pollinators are visually percep- talk along with illuminating broadscale mits are bees shielded from these negative tive of ultraviolet (UV) color, flower geographic patterns in bee specialization, effects of warming. Over a 500 m altitu- Continued on page 16 Page 15 both direct measures of pollen transfer efficiency and a correlated trait (pollen-to- ovule ratios) in a phylogenetic context, I show that pollinia evolved in an ancestral flower that already had functionally aggre- gated and highly efficient pollen transfer. However, power analysis indicates that the sample size is not yet adequate to detect if Apocynaceae with pollinia (milkweeds) have significantly more efficient pollen transfer than those without. Comparative climate niche analysis links the origin of pollinia to ecological conditions generated by the aridification of Africa during the Cenozoic, consistent with the hypothesis that ecological adversity has selected for the evolution of efficiency.
Nathan Muchhala University of Missouri – St. Louis Sam Droege (USGS Patuxent Wildlife Research Center) speaks about North “Bats, birds, and bellflowers: The evo- American pollen-specialized bees. (photo by Ken Wurdack) lution of specialized pollination systems in the Neotropics” Abstracts UV pigmentation. Instead, I present data Animal pollination is thought to have Continued from page 15 that suggest how selection via abiotic fac- tors (in conjunction with pollinators) may played a central role in angiosperm diver- color patterns in the UV spectrum have drive variation in UV floral pattern. sification, especially in the tropics, where particularly sparked interest in pollination over 98% of plants are animal-pollinated. biologists. A common UV ‘bullseye’ floral Research in my lab combines experi- Tatyana Livshultz phenotype has long been held to function ments and theory to explore the ecology Drexel University as a nectar guide for pollinators, increas- and evolution of plant-pollinator interac- ing pollination efficiency. I experimentally “Putting milkweeds in context: The tions, with a focus on bat pollination in test how variation in floral UV pigmenta- evolution and function of pollen aggrega- the Neotropics. In this talk I begin by tion phenotype affects pollinator behavior tion in Apocynaceae” discussing research on an unusual species using phenotypic manipulation, and then Pollination is an inefficient process. of nectar bat which can launch its tongue address whether pollinators may contrib- On average, less than 1% of exported 1.5 times its body length, an extension ute to UV floral variation in nature using pollen grains are deposited on conspe- more than double that of other bats and the common plant, Argentina anserina cific stigmas. Pollen transfer is an order longer than any other mammal. Unique (Rosaceae). Pollinators preferentially of magnitude more efficient (ca. 25% on adaptations allow it to store its tongue in visit flowers with a distinct UV bullseye average) in species with pollen packaged its rib cage. Experiments suggest that this pattern, however UV pattern variation into pollinia, yet pollinia have evolved bat is involved in a coevolutionary race does not affect their alighting, or foraging in only two families, Orchidaceae and with long-tubed flowers; tongue elonga- behavior. There is pronounced variation in Apocynaceae, indicating strong fitness tion allows bats to reach more nectar, the size of the UV bullseye in nature, and costs under most ecological conditions. I while flower elongation maximizes pollen geographically shifting pollinator prefer- develop the hypothesis that pollinia and transfer. In the second part of the talk, I ence can contribute to patterns of floral elevated pollen transfer efficiency are present evidence for two cases of character variation across space. While UV pigmen- adaptations to conditions of strong mate- displacement in response to competition tation patterns are variable within species, finding Allee effects and high levels of for pollination. The first involves overdis- they also differ substantially among taxa, interspecific competition for pollination, persion in flower morphology: co-occur- potentially owing to pollinator-mediated allowing plants to continue outbreed- ring Burmeistera were found to place their trait divergence. I test whether pollinators ing, but at the cost of drastic reduction pollen in different regions of bats heads’, may drive phenotypic variation among in the number of mates. Apocynaceae, thus maximizing conspecific pollen trans- taxa in the Potentilleae (Rosaceae) tribe with ca. 5000 species, have a diversity fer despite sharing bats as pollinators. The using a biogeographically-informed sister of floral morphologies and degrees of second involves overdispersion in flower taxon analysis. Interestingly, species that pollen aggregation, including monads, color: co-occurring Iochroma were found co-occur have more similar UV pigmenta- tetrads, and pollinia, making them an ideal to differ significantly in color, as perceived tion phenotypes than those that grow in lineage to investigate the evolution of by their primary pollinators (hum- spatially distinct regions, suggesting a lack pollen aggregation and its adaptive value mingbirds and bees), relative to random of reproductive character displacement for using the comparative method. By placing expectations. I end the talk with a brief
Page 16 overview of other ongoing projects in the generalized pollinator hubs such as Cir- tion with the invader. We also witnessed lab. Together, this research shows some of sium arvense and Achillea millefolium. In subsequent growth of tree populations the ways that coevolution and competition addition, I will explore our current knowl- pollinated by the honeybees, and a boom have shaped the highly specialized pol- edge of geographic variation in floral scent in native bee resources that were offered lination systems found in the tropics. - the potential for local scent dialects - in via honeybee pollination services. Some the context of the Geographic Mosaic solitary bees, on the other hand, did not Robert Raguso Theory of Coevolution, with reference to respond to the Africanized bee invasion, Cornell University an ongoing study on the genus Oenothera. but maintained their broad relationship with flowering plants that sustain them, “Floral scent: The dark matter of plant- in remarkably flexible fashion, and in the pollinator interactions” David W. Roubik Smithsonian Tropical Research face of long-term climate changes and Flowers are not merely objects of Institute, Panama hurricanes. Our primary conclusion is that aesthetic beauty; they serve as engines of the wide diversity in native vegetation, “Whose bees are these? The pollen tax- biological diversity, lynchpins of ecologi- particularly in the tropics (and also in the onomy of bee nests and its story in recent cal stability and fonts of human ecosystem Washington, DC area), insures that flower decades” services from agriculture and floriculture visitors can adjust to and survive the tests to cosmetics. Botanists have long been We need to know more about why the of floral resource availability and other dazzled by the visual aspects of floral populations of our bees go up and down, challenges. display (color, shape and pattern). In con- and a good deal more about what makes trast, the chemical aspects of floral func- those bees in the first place. Making a tion, from the scents that guide pollinator pollen reference collection to a particular attraction and learning to the composition flora is both time-consuming and reward- of the nectars, resins and oils that reward ing. For bee ecology, there is no greater their visits, somehow remain peripheral source of detailed information. We have to the central bodies of ecological and characterized honey origins from its pollen evolutionary theory concerning pollina- content, and pollen in orchid bee and soli- tion. However, recent technological and tary bee nests—without barcoding—but conceptual advances have made it easier with replicated work. Our studies can, in to analyze and manipulate floral chemistry, contrast to all but a very small number of Symposium particularly floral scent, and a growing places that have established molecular ref- Continued from page 1 body of evidence points to more central erence libraries to the local flora, usually creativity and curiosity. His research on roles for scent in mediating floral isola- identify pollen grains to species. Armed Barro Colorado Island, a 1,560-hectare tion, constancy, gene flow and defense. with this insight, we can begin to picture island in the Panama Canal, makes use of I will highlight several recent manipula- the consumer’s dilemma (for a bee) and botanical knowledge, reference collec- tive studies, in which visual signals were plant community efforts to obtain pollina- tions, photographs, and field and labora- tracked and controlled, to illustrate the tion service and reproduce. tory work. Among his studies are orchid unexpected roles played by scent in oth- An early question that motivated our bees, honey bees, Africanized honey bees, erwise well-studied model systems. Floral building of two reference pollen libraries, Centris bees, and megachilids. scent provides critical mechanisms that one for Panama and one for the Yucatán Roubik’s pollen research has also explain conditional reproductive isolation Peninsula of Mexico, was the escape included studies of honey bee competi- among sympatric Ipomopsis aggregata and explosive range extension of African tion with native bees. A 17-year study and I. tenuituba, balance floral defense honeybees in the Neotropics. What do from the Yucatan Peninsula in Mexico and pollinator-mediated selection on floral they eat, and which plants are primar- shows the impact of invasive Africanized form in Polemonium viscosum, and dictate ily responsible for sustaining them? By honey bees on solitary megachilid bees. the network structure of floral visitors to quantifying their pollen taken to hives, and He found honey bees may have forced the also pollen ingredients of their honey, we native bees to switch their food plants. Sponsors of the saw striking evidence that many primary Pollen from plants in the families Euphor- forest resources of the invasive honeybees th biaceae and Anacardiaceae were present 14 Smithsonian are trees, and often not pollinated by bees, in lower amounts in megachilid nests after Botanical Symposium but yielding huge surpluses of nectar and honey bee invasions, while Sapotaceae, pollen for “cleanup” organisms, such as Fabaceae, and Rubiaceae pollen were • Cuatrecasas Family Foundation social bees and their colonies. Much of present in great amounts in the same nests. • National Museum of Natural their food was “leftovers” from nocturnal Tatyana Livshultz (Drexel University) History flowers designed to feed and attract larger continued the focus on pollen with her • Department of Botany animals, like bats or moths. We docu- talk, “Putting milkweeds in context: The • Office of the Associate Director mented the decline of native bee resource evolution and function of pollen aggrega- for Science use when displaced by the invasive hon- tion in Apocynaceae.” Livshultz studies eybee, but saw rapid ‘resource partition- the structural and functional diversity of • United States Botanic Garden ing’, as local bees escaped food competi- Continued on page 18 Page 17 Symposium may have created the selective pressure for tribute to UV variation. He showed that Continued from page 17 evolution of increased PTE and pollinaria insects are more attracted to UV bullseye in milkweeds. in Potentilla anserina, that the UV pattern, pollen. The most common type of pollen The next two speakers spoke about not the reflection, increases attractiveness, aggregation are tetrads, 4 pollen grains unseen floral attractants—scent and and that the bullseye does not serve as a dispersed as a unit. Orchidaceae and ultraviolet pigmentation. First was Robert nectar guide. Apocynaceae have pollinaria, two or more Raguso (Cornell University) who dis- The pollinator communities of Poten- pollinia (entire pollen content of an anther cussed “Floral scent: The dark matter of tilla anserina change with altitude (bees sac fused into a mass) attached at a trans- plant-pollinator interactions.” Raguso predominantly at lower altitudes and flies lator. Livshultz showed that aggregated explained that in studies of the structure at higher altitudes), as does the bullseye pollen and translators evolved three times of plant-pollinator networks, floral scent is size (larger bullseye at higher altitudes). independently within Apocynaceae. almost never mentioned, yet it can be just Koski found that by manipulating the size Livshultz demonstrated how aggrega- as important as visual and tactile patterns. of the bullseye at low and high altitude tion affects pollen performance through He showed how scent can affect seed fit- sites, both bee and fly preference for bulls- measures of pollen transfer efficiency ness and structure floral networks through eye shifted with altitude. Plants with larger (PTE), the fraction of exported pollen attraction and repellence. In one study bullseyes received more visits and pollen grains that reach conspecific stigmas. She floral fragrances ofCirsium arvense and by both pollinators at higher altitudes, discussed how milkweed pollinarium Achillea millefolium were swapped, and while plants with smaller ones received evolved in an ancestor that already had the links between insect visitors to the two more pollen at lower sites. In terms functionally aggregated pollen and high plant species were restructured until the of abiotic factors, he found that larger PTE. The evolution of the pollinarium scents dissipated. In Penstemon digitalis, bullseyes protect pollen from UV-damage. further elevated PTE. She explained that where the corolla limb is scentless and the Looking at macroevolutionary patterns of current evidence does not indicate that nectar tube has volatiles, Raguso showed diversity in the genus Potentilla, Koski evolution of the gynostegium nor tetrads how floral scent emission and floral explained that the UV bullseye is larger and translators in Apocynum and Periplo- display were under selection, and flower in more alpine species which experience coideae elevated PTE. Finally, Livshultz size and color were not. This suggests that higher UV-irradiance and cooler tempera- put milkweeds in a paleoecological con- smelling stronger benefits reproductive tures. He concluded that while pollinators text and explained that drought-induced success in this flowering species. are important in driving variation among mortality of plants and pollinators during Raguso also explained how scent can populations of Potentilla, it is the abiotic early Oligocene aridification of Africa help integrate complex floral phenotypes factors that drive diversity at both micro- and mediate reproductive isolation through and macroevolutionary levels. Acknowledgements attraction, preference and constancy. Moving the discussion from insect to Dracula lafleurii, an orchid species from vertebrates, Nathan Muchhala (University The success of the Symposium was Ecuador, is pollinated by flies that breed of Missouri-St. Louis) discussed “Bats, due to the significant time and efforts in mushrooms, and Dracula flower parts birds, and bellflowers: The evolution of of the following people: resemble mushrooms. In a study using specialized pollination systems in the Neo- 3d-printed silicon models of Dracula tropics.” To examine bat-flower coevolu- Organizers flowers, Raguso was able to tease apart tion, Muchhala examined the efficiency • Laurence J. Dorr, visual, olfactory, and tactile responses in of bat tongue at extracting nectar and • Seán Brady which a combination of the three gets the the ability of bats to transfer pollen. He • Gary Krupnick best response by flower visitors. A study showed how tongue length varies among • Sue Lutz examining visitation by Hyles lineata three bat species, with Anoura caudifer • Ari Novy hawkmoths to scent-augmented and color- and A. geoffroyi having short tongues (3.7 • Sylvia Orli swapped Ipomopsis flowers demonstrated cm and 3.9 cm, respectively) while A. • Susan Pell that moths require both scent and high fistulata have long tongues (8.5 cm). The • Eric Schuettpelz visual contrast to feed on flowers at dusk. diet of Anoura fistulata includes many • Holly Shimizu He concluded by stressing the importance plant species, but not the short corolla • Warren L. Wagner of scent combined with visual traits in pol- Burmeistera. In pollen transfer experi- • Kenneth Wurdack lination studies. ments in the field, Muchhala showed how • Elizabeth Zimmer Matthew Koski (University of Vir- angle and depth are more important than Support ginia) followed with a talk on “The duration and force –bats lift their heads for • Mary Ann Apicelli evolutionary ecology of ultraviolet floral deeper feeding which led to greater trans- • Carol Youmans pigmentation at micro- and macroevolu- fer of pollen. He argued that Anoura fistu- tionary scales.” Ultraviolet (UV) pigmen- lata appears to have evolved its incredibly Photographer tation leading to a bullseye pattern has long tongue through coevolutionary races • Kenneth Wurdack been found in at least 23 of 62 angiosperm with long-tubed flowers. And many others who had helped in a orders. Koski’s research focused on how Muchhala also spoke on interspecific myriad number of ways. UV bullseye mediates pollinator behavior competition (plants competing for pol- and if pollinators or abiotic factors con- linators), floral character displacement,
Page 18 and evolution. He discussed his research on overdispersion of color in Iochroma, a hummingbird- and bee-pollinated genus of 33 species of shrubs and small trees throughout the Andes. Looking at color differences, phylogenetic distances, and differences in plant morphology (lengths of petiole, leaf, and pedicel), Muchhala asked whether coexisting communities of species differ more than expected by chance. He found that hummingbird and bee pollinators are driving the difference in color among the species, and that there is ongoing selective pressure among the plant species to be different in color as much as possible to reduce the costs of receiving pollen from close relatives. The talks concluded with a presenta- tion by Candace Galen (University of Missouri) who spoke about “Shifting baselines and changing partners: Ecologi- Poster presenters and guests interact at the opening reception of the Smithsonian cal and evolutionary responses to climate Botanical Symposium. (photo by Ken Wurdack) change in alpine bumble bees and their st host plants.” To examine the impacts of The Symposium ended with a recep- Taxonomy in the 21 Century” (2001); global climate change on plant-pollinator tion and poster session in the Conserva- “The Convention on Biological Diversity” relationships, Galen conducted field stud- tory of the United States Botanic Garden. (2002); “Botanical Frontiers in South- ies at three sites in the Colorado Rocky A group of 14 presenters displayed their east Asia” (2003); “Botanical Progress, Mountains where overnight lows are on posters and spoke about their research Horticultural Innovations, and Cultural average 2oC warmer and flower production ranging in topics from floral rewards and Changes” (2004); “The Future of Floras: has seen a 60 percent decrease since the the diet of hummingbirds and bats to New Frameworks, New Technologies, 1970s. Galen examined whether Bombus honey bee health and the conservation of New Uses” (2005); “Island Archipelagos: species are broadening their food choices. endangered species. Cauldrons of Evolution” (2006); “Partners th She found that the foraging breath of Next year’s 15 Smithsonian Botani- in Evolution: Interactions, Adaptations, bumble bees has more than doubled, and cal Symposium will take place on Friday, and Speciation” (2008); “Genes, Genom- that flower shape of host plants has also May 21, 2017, with a theme yet to be ics and Genome Evolution in Plants” st broadened—many short-tubed flowers are determined. Be sure to check the sympo- (2009); “Food for Thought: 21 Century now among those they visit. sium website at
Page 19 Art by Alice Tangerini
Oenothera macrocarpa Nutt. At the 2016 Smithsonian Botanical Symposium, speaker Sam Droege (U.S. Geological Survey) talked about pollen specialist bees which have evolved obligate associations with flowering host plant genera. Oenothera is a recurrent host plant genus associated with specialist bee species in the United States. Oenothera macrocarpa is a known host plant for the sweat bee, Lasioglossum oenotherae. Alice Tangerini illustrated O. macrocarpa for a T-shirt design for the Botany 2011 conference in St. Louis, Missouri. The type locality for this species is along bluffs along Meramec River near the Shaw Nature Reserve of the Missouri Botanical Garden. This species was selected for the shirt since the meetings were in St. Louis, and Missouri Botanical Garden director Peter Raven and his students worked on Onagraceae. The drawing began as a pen and ink illustration based on US herbarium specimens and field photographs. Digital color was added for the shirt logo.
Department of Botany PO Box 37012 NMNH, MRC-166 Washington DC 20013-7012
Official Business Penalty for Private Use $300
Page 20