An updated description of the collections and history of The New York Botanical Garden Herbarium (NY): 1995–2015 MATTHEW C. PACE, NICOLE TARNOWSKY, ELLEN D. BLOCH, AMY WEISS, CHARLES ZIMMERMAN, AND BARBARA M. THIERS William and Lynda Steere Herbarium, New York Botanical Garden, 2900 Southern Blvd., Bronx, NY 10458-5126, USA; e-mail: [email protected]

Abstract. In celebration of The New York Botanical Garden’s 125th anniversary, we present an updated description of the specimen holdings and activities of The William and Lynda Steere Herbarium of The New York Botanical Garden (NY) from 1995 to 2015. During this time, the collection grew to approximately 7.8 million specimens, a new International Plant Sciences Center was built to house the Herbarium and LuEsther T. Mertz Library, many plant families and other taxonomic groups grew in scope and taxonomic comprehensiveness, Southeast Asia developed as a new focus of collecting activities, and the Herbarium emerged as a leader in specimen digitization. Key Words: Biodiversity, collections management, herbarium, natural history collection, North American flora, South American flora.

The period of 1995–2015 has been one of tremendous change and opportunity for The William and Lynda Steere Herbarium (NY) of The New York Botanical Garden. We moved the Herbarium to new quarters (Fig. 1) and have built a completely new collection of digital representations of the physical specimens. The creation, growth and maintenance of this new virtual collection require as much attention as the physical collection (Thiers et al., 2016, this issue). Recently growing to approximately 7.8 million specimens, the Steere Herbarium remains the largest herbarium in the Western Hemisphere, and is the second largest such collection in the world (Thiers, continuously updated). The objectives of the Steere Herbarium remain the same as they have for the 125-year history of the institution: to document the scientific research of the Garden’s staff, and to represent through specimens the biodiversity of the algae, bryophytes, fungi, lichens, and vascular plants of the world, with special emphasis on the Americas. Serving the research community beyond Garden staff has been equally important, and the primary impetus for the specimen digitization program was to increase access to our holdings, which continue to be available through loan and visitation. Because of two economic crises, a trend in universities to move away from organismal research, and increasing restrictions and regulation of

international shipment of biological material, many natural collections have reduced their programs. However, the Herbarium has continued to expand and improve the collection, due to the vision, tenacity, and dedication of those who managed it during its first century, and it is still one of the most valuable resources in the world for biodiversity research. The purpose of this article is to update the article on the history of the Herbarium by Holmgren et al. (1996), by summarizing the changes that have taken place in the acquisition, maintenance, curation of the specimens, and the data sharing of the Herbarium over the past 20 years. Accessions The past 20 years have been a period of expansive growth for the Steere Herbarium (Table I; Fig. 2), which now houses approximately 7.8 million specimens, composed of approximately 2 million algae, bryophytes (mosses and hepatics), fungi, and lichens, and 5.8 million vascular plants (ferns and lycophytes, gymnosperms, and angiosperms). This total number includes ca. 50,000 non-vascular plant and fungal type specimens and 98,700 vascular plant type specimens. An average of about 52,000 newly mounted specimens were added to the collection every year,

Brittonia, DOI 10.1007/s12228-016-9422-8 ISSN: 0007-196X (print) ISSN: 1938-436X (electronic) © 2016, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

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FIG. 1. A. The International Plant Sciences Center, home of the Steere Herbarium and the LuEsther T. Mertz Library. The building was designed with few windows so as to minimize damage to the specimens from sunlight, dust, or possible pests. B. Curation and research activities within the Herbarium.

making the Steere Herbarium one of the most active herbaria in the world. Accession of TABLE I Number of added specimens, 1995–2015, including staff collections and orphaned herbaria.

Group Bryophytes Fungi Lichens Algae Vascular Plants Total

Number of specimens added 91,969 95,682 148,244 94,948 703,340 1,043,576

collections continued to focus on the Steere Herbarium’s historic areas of strength, including the northern Andes, Brazil, the Caribbean, the Guiana Highlands, and North America, while expanding to new areas of interest, including Southeast Asia, Australia, and Oceania (Appendix 1). Specimen accessions by division are reflected in Fig. 2, where for the first time the number of annual accessions of non-vascular plants and fungi recently surpassed those for vascular plants, primarily due to the acquisition of several orphaned herbaria. In total, the Steere Herbarium grew by 1,043,576 specimens from 1995 to 2015 (Table I). Additionally, in 2015 the entire Brooklyn Botanic Garden herbarium (BKL) was transferred to the

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FIG. 2. Annual newly accessioned specimens in the Herbarium (based on mounting records in the vascular plant herbarium and incoming shipments to the non-vascular plant and fungal herbarium), 1995–2015.

Steere Herbarium as a long-term loan, where it will remain until the science program is reestablished at Brooklyn, projected to be some time in the next ten years. The Brooklyn herbarium remains separate from the general collection of the Steere Herbarium and is open for research. NON-VASCULAR PLANT AND FUNGAL COLLECTIONS

Non-vascular plant and fungal collections made by Herbarium research staff primarily focused on the United States of America, Chile, Australia, Costa Rica, and Canada (see also Daly, 2016, this issue). The collections of James Lendemer significantly increased the number of lichen specimens housed in the Steere Herbarium, with 37,824 databased specimens added between 2001 and 2015 (see also Lendemer & Harris, 2016, this issue). The adoption of a growing number of orphaned herbaria has contributed significantly to increasing the size and scope of the Steere Herbarium’s non-vascular plant and fungal collection. Although this has been a scientific boon for the Herbarium, increasing the tremendous taxonomic and geographic depth and breadth of its holdings while complementing ongoing staff collections, it also reflects national declines in funding and general support for herbaria and other natural history collections. Between 1995 and 2015, the Steere Herbarium accepted approximately 246,000 non-vascular plant and fungal collections from 15 institutions or

individuals (Appendix 2). In 2002, the Steere Herbarium diatom collection was donated to the California Academy of Sciences (CAS). VASCULAR PLANT COLLECTIONS

Vascular plant collections made by the Garden research staff primarily focused on the countries of Brazil, the United States of America, Colombia, Bolivia, and Cuba (Appendix 1), and more generally on the Amazonian and Atlantic Coastal regions of Brazil, the northern Andes, Intermountain and Eastern North America, and Southeast Asia, reflecting the interests of staff researchers. Families with significant accession increases include Anacardiaceae, Burseraceae, Cyperaceae, Ericaceae, Lecythidaceae, Melastomataceae, Scophulariaceae s.l., and Solanaceae. Additionally, the Steere Herbarium received many significant specimen gifts, including thousands of specimens from John L. Clark (Ecuador), Wayne Longbottom (Mid-Atlantic U.S.A.), George Schaller (Tibetan Plateau), Jerry Tiehm (Intermountain U.S.A.), and David C. Thornburg (Southwestern U.S.A.). In contrast to the non-vascular plant and fungal collection, only a few orphaned vascular plant herbaria were incorporated during this time (Appendix 2). Physical upgrades and infrastructure One of the signature achievements for The New York Botanical Garden and the Herbarium

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Directorship of Patricia Holmgren was the conception, design, and building of the International Plant Science Center (Fig. 1), which was added to the northwest corner of the original Beaux-Arts Museum and Library building, to house The William and Lynda Steere Herbarium, The LuEsther T. Mertz Library, and nine study rooms for visiting researchers. Planning for this state of the art building began in 1996, with construction commencing in 1998. From 2000–2001, the massive and complicated task of moving the entire herbarium into this new building was planned and overseen by Jacquelyn Kallunki, who coordinated a team of 58 Garden staff, interns, and volunteers for a total of about 3300 hours. The new five-floor International Plant Science Center was specifically designed to meet the particular needs of herbarium specimens, including controlled environmental conditions, an emergency sprinkler system, and more than 150 Spacesaver compact mobile shelving units with dozens of new water and insect-proof cabinets, allowing for better curation and conservation of the specimens, as well as providing room for visiting researchers and the continued expansion of the collection. Funding was generously provided by the United States Congress—with leadership from Senators Daniel Patrick Moynihan and Alfonse M. D'Amato—through the United States Department of Agriculture, the LuEsther T. Mertz Charitable Trust, Edward P. Bass of the Philecology Trust, Shelby White and Leon Levy, Arthur and Janet Ross, The Starr Foundation, the National Science Foundation (DEB-9710105; supplement to DBI-9808824), The Prospect Hill Foundation, The Bank of New York, J. P. Morgan and Co., Incorporated, The City of New York, and New York State. Additional grants from the National Science Foundation afforded the purchase of microscopes and computers for the International Plant Studies Center. In 2002, the William and Lynda Steere Herbarium and the LuEsther T. Mertz Library officially opened for research. Once the herbarium move was completed, attention turned to renovating the old Museum and Library Building. In 2006, a beautiful new mounting room (Fig. 3) and Digital Imaging Center were created on the 4th floor; previously, the mounting room was located on the 1st floor of the Museum and Library Building, connected to the sorting room. The new mounting and imaging facilities are now part of regular herbarium tours to visiting school groups, providing a holistic view of the modern herbarium. The creation of

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the Digital Imaging Center also signaled the Herbarium’s pioneering initiative to create a publicly accessible digital database of its entire authoritative collection: the C. V. Starr Virtual Herbarium (Thiers et al., 2016). This process, with the goal of providing full specimen data and an image for all of the Herbarium’s specimens, began in 1996 and has been supported by a dozen National Science Foundation grants, focusing on all plant groups and on several geographic areas. In 2014, a major grant from the National Science Foundation provided funds for the installation of new extra tall cabinets in a portion of the 1st floor of the herbarium, to allow expansion and better curation of the palms, lichens, and algae. Collections management and curation The Steere Herbarium, through the efforts of staff researchers in the Herbarium, Institute of Systematic Botany, and The Lewis B. and Dorothy Cullman Program for Molecular Systematics, remains one of the most authoritatively curated herbaria in the world. From 1995 to 2015, families and groups that received particularly thorough curation, and which acquired many new specimens, include Agaricales and Boletales (Roy Halling), Arailiaceae (Gregory Plunkett), Arecaeae (Andrew Henderson), Aristolochiaceae (Lawrence Kelly), Burseraceae (Douglas Daly), Characeae (Kenneth Karol), Cyperaceae (Robert Naczi, W. Wayt Thomas), Ericaceae (James Luteyn, Paola Pedraza), Fabaceae (Rupert Barneby, Benjamin Torke), Lecythidaceae (Scott Mori), Lichens (James Lendemer), Melastomataceae (Fabian Michelangeli), Pleurocarpus mosses (William Buck), Pteridophytes (John Mickel, Robbin Moran), Orchidaceae (Matthew Pace), Rubiaceae (Brian Boom), Rutaceae (Jacquelyn Kallunki), Scrophulariaceae s.l. (Noel Holmgren), Solanaceae (Michael Nee), Symplocaceae (Lawrence Kelly), and Xyridaceae (Lisa Campbell). The ethnobotanical collections also grew due to the expansive research of Michael Balick and Ina Vandebroek. Curation of various families was often facilitated and complemented by National Science Foundation grants focused on digitization of various groups and/or geographic regions (e.g., flora of eastern Brazil; flora of the Caribbean; PBI: Miconieae [Melastomataceae]) (Thiers et al., 2016, this issue). The Herbarium’s Cold Storage Room contains approximately 50,000 unidentified and unmounted specimens (2000 bundles) collected by past and

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FIG. 3. The current spacious, functional, and beautiful mounting room, on the 4th floor of the Beaux-Arts Library Building.

current staff researchers. These specimens belong to taxonomic groups for which there are no current specialists or for which identifications have not yet been received from existing specialists; some of the specimens have remained in the Cold Storage Room for the last 40 years awaiting an authoritative identification. We thoroughly renovated the Cold Storage Room and inventoried, reorganized, and fumigated these collections from 2007 to 2008. The inventory facilitated the conservation of these valuable specimens—many of which likely represent new as-of-yet undescribed species (Bebber et al., 2010)—and the development of a strategy to obtain identifications. Over the next few years we will begin to process, distribute, database, and image these undetermined specimens with the hope of gaining full identifications. In 2000, Patricia Holmgren retired as Director of the Herbarium with achievements including the building of the International Plant Science Center, initiating efforts to digitize the collection, and editing Index Herbariorum while transitioning it to a digital platform. Holmgren continues to serve the Herbarium as Director Emerita. Barbara Thiers, previously Associate Director of the Herbarium, was appointed as the new Director of the Herbarium, leading the herbarium as it became a pioneer in specimen digitization and data sharing, and in 2014 concurrently became Vice President for Science. The years 2014 and 2015 saw the retirement

of three NY administrative giants, who together represented over 70 years of institutional knowledge: Jacquelyn Kallunki (Associate Director of the Herbarium), Thomas Zanoni (Senior Collections Manager), and Stella Sylva (Collections Manager). Kallunki, a world authority on general Neotropical plant identification, Goodyera (Orchidaceae), and Rutaceae, continues her systematic research on Rutaceae as Curator Emeritus. Outreach From 1995 to 2015, the Steere Herbarium developed into a leader in the worldwide effort to digitize natural history collections; we share our growing collections and updated resources with the botanical community at every opportunity, impacting researchers, students, and members of the general public (Fig. 4). Our citizen science and volunteer programs have expanded in the past 20 years to include participation in many aspects of herbarium work: our volunteers donate their time to mount and image specimens, as well as transcribe specimen data in the online portal, Symbiota. In 2015 alone, more than 20 volunteers captured images of approximately 25,000 specimens and mounted 6500 vascular plant specimens. Making the information from specimens accessible to the widest possible audience continues to be a major goal of the Herbarium. Access has been facilitated

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over the past 20 years by capturing data and images for over two million of our specimens and providing this online to the public, via the C. V. Starr Virtual Herbarium. Building on this data, we have developed eFlora and eMonograph pages for many of our scientists. Online publications include Flora of French Guiana, Flora of the Osa Peninsula, The Lichens of the Ozarks, Lichens of the South Eastern Plains, the Elaphoglossum and Megalastrum Pages (ferns), Lecythidaceae pages, the Miconiae pages, and the Barneby Legume Catalogue. Providing access to our physical collection for both the scientific community and the general public remains a priority. Annually on average, about 200 researchers spend a collective ca. 2000 days using the herbarium. We send an annual average of 17,000 specimens on loan to other institutions, underscoring the critical role the Steere Herbarium collections play in modern systematics, genomics, conservation biology, ecology, ethnobotany, and structural botany. Our first large-scale open house to the general public was provided in the spring of 2002 to celebrate the opening of the International Plant Science Center. Since 2013 we have annually opened our doors in subsequent open houses, giving insight into our

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collections, best practices, and overall Science program. Herbarium staff provide an average of 35 private tours per year, many of which are for high school and university classes (Fig. 4). By emphasizing the pivotal role that herbarium collections play in understanding global and regional biodiversity, the consequences of climate change, invasions of weeds and plant pests, how plants create the habitats that support animals, etc., we hope to inspire future generations towards careers in botany. The Steere Herbarium has been able to reach the public even further through several media outlets. Between 2012 and 2015, four separate films featuring the herbarium were made: PBS Treasures of New York: The New York Botanical Garden (aired late June, 2013); an independent film by Alister Sanderson entitled BHerbarium: Inside The New York Botanical Garden^ (http://vimeo.com/ 69637924); a promotional video for the Global Plants Initiative by JSTOR (http://vimeo.com/ 74645215); and an article about the World Flora Online project for the BGreat Big Story^ series on CNN online (http://www.greatbigstory.com/stories/ photographing-every-plant-in-the-world). Steere Herbarium specimens have also been featured in

FIG. 4. Tour of the Steere Herbarium to visiting university students, led by Nicole Tarnowsky, Assistant Director of the Herbarium. The specimen held by Tarnowsky is of Chiliotrichum diffusum (G. Forst.) Kuntze (Asteraceae), collected from Tierra del Fuego during the circumnavigation of the world by Captain J. Cook, J. Banks & D. C. Solander s.n., 15 Jan 1769–21 Jan 1769 (NY barcode 22391).

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various other social media, including our NYBG blog, ScienceTalk (http://blogs.nybg.org/sciencetalk/), Facebook page (https://www.facebook.com/ NYBGHerbarium) and Instagram account (https:// w w w. i n s t a g r a m . c o m / n y b g h e r b a r i u m / ; #nybgherbarium).

Thiers, B. Continuously updated. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden's Virtual Herbarium (http:// sweetgum.nybg.org/science/ih/) ———, M. C. Tulig & K. A. Watson. 2016, this issue. Digitization of The New York Botanical Garden Herbarium. Brittonia 68(3).

Future directions The next 20 years offer tremendous opportunity for the Steere Herbarium. Specimen holdings will continue to grow in our established strengths of North and South America and the Caribbean, while concurrently growing in developing strengths: lichen diversity, Orchidaceae, the flora of the New York City Metro area, and the flora of Myanmar, Vietnam, Malaysia, and Oceania. Continued digitization efforts and sharing of data will aim to provide digital representations for all specimens within the next 20 years. Building on the dedication and curatorial and scientific excellence of our predecessors—including Nathaniel Lord Britton, Henry Gleason, Bassett Maguire, and Patricia Holmgren—the Steere Herbarium continues to be one of the finest such collections in the world, and a critical source of knowledge for the world’s plants, fungi, algae, and lichens.

Appendix 1 Origins of staff collections, 1995–2015; top 30 countries only. Countries with asterisks are the ten largest sources of new staff collections during this period. Australia * Belize Bolivia * Brazil * Canada Chile Colombia * Costa Rica * Cuba * Dominican Republic Ecuador * Federated States of Micronesia French Guiana Guatemala Guyana

Malaysia Mexico Myanmar New Zealand Palau Panama Papua New Guinea Peru * Puerto Rico St. Eustatius Sweden United States of America (excluding territories) * Vanuatu Venezuela * Vietnam

Acknowledgments The authors wish to thank the many staff members from 1995 to 2015 who compiled in Annual Reports the raw data used in this manuscript; Patricia Holmgren, Jacquelyn Kallunki, and Lawrence Kelly for helpful comments during review of this manuscript; the Board of The New York Botanical Garden for their support.

Literature cited Bebber, D. P., M. A. Carine, J. R. I. Wood, A. H. Wortley, D. J. Harris, G. T. Prance, G. Davidse, J. Paige, T. D. Pennington, N. K. B. Robson, R. W. Scotland & D. B. Wake. 2010. Herbaria are a major frontier for species discovery. Proceedings of the National Academy of Sciences of the United States of America, 107(51): 22169–22171. Daly, D. 2016, this issue. Around the lunch table: Systematics and systematists at The New York Botanical Garden. Brittonia 68(3). Holmgren, P., J. Kallunki & B. Thiers. 1996. A short description of the collections of The New York Botanical Garden Herbarium (NY). Brittonia 48: 285–296. Lendemer, J. C. & R.C. Harris. 2016, this issue. The NYBG Lichen Herbarium: a unique resource for fungal biodiversity research and education. Brittonia 68(3).

Appendix 2 Collections of algae, bryophytes, fungi, lichens, and vascular plants donated to NY, 1995–2015. Biechele, L.: ca. 400 lichens and bryophytes of Delmarva Peninsula (2010) Dey, J.: 35,000 lichens from the southeastern U.S. (2012) California Academy of Sciences (CAS): ca. 3000 fungi (2002) Field Museum of Natural History (F): ca. 90,000 algae (2015) Kansas State University (KSC): ca. 17,000 fungi (1997) Kohlmeyer, J.: 20,000 marine fungi (2009) Lay, E.: ca. 17,000 lichens of eastern U.S. (2010) Mianus River Gorge herbarium: 246 vascular plants (1996) Miller, N. G.: ca. 5000 bryophytes (2011) Herb Society of America: 516 vascular plants (1996) Pennsylvania State University at Mont Alta (PACMA): ca. 3700 fungi (2015) Reed, C.L.: 9238 lichens (2001) Tulane University (NO): 10,000 fungi (2000) University of Rhode Island (KIRI): 2111 fungi (1995) University Tennessee (TENN): ca. 5000 bryophytes & lichens (2015) Virginia Polytechnic Institute (VPI): 25,000 fungi (2012) Whittier, H. O.: ca. 5000 bryophytes mostly from the South Pacific (2010)