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Department of Vertebrate Zoology

The mission of the Department of Vertebrate Zoology is to discover, describe and classify the world’s species of vertebrates and interpret the evolutionary history of this high profile diversity to meet the needs of science and society.

Research in the Department of Vertebrate Zoology is organized into four divisions: Amphibians and Reptiles, Birds, Fishes, and Mammals. Research studies extend across the spectrum of systematics, morphology, molecular biology, biogeography, life history, behavior, and ecology of fishes, amphibians, reptiles, birds, and mammals with strengths in phylogeny and revisionary studies within these groups. Geographical areas of particular research interest include North, Central and South America; Africa; and the Indo-Pacific region and adjoining areas in southern Asia.

Worldwide collections of preserved specimens and extensive osteological collections are the basis for monographic studies of vertebrate species and their higher taxa, and for related studies focused on the evolution and ecology of vertebrates. The vertebrate collections trace their origin to the two boxcars of specimens that Spencer Fullerton Baird, one of the first Secretaries of the Smithsonian, brought with him in 1850. Since that time, the Department of Vertebrate Zoology has grown with responsibility to maintain the foremost international collections of vertebrate animals, comprising the world’s largest collections of fishes (approximately 6 million specimens), mammals (590,000 specimens), and amphibians and reptiles (570,000 specimens), plus the world’s third largest collection of birds (600,000 specimens). The research value of each Division’s holdings is amplified by many historically important series including 15,803 primary type specimens. Accordingly, the department is recognized internationally for the systematic and geographic comprehensiveness of its collections and for its influential, high profile research programs in systematic biology and associated fields.

Division of Amphibians and Reptiles
Research in the division covers a wide spectrum of biological topics and geographic areas. Most research is collections based and emphasizes the evolution, biogeography and systematics of selected groups of frogs, lizards, snakes and turtles from North America, tropical America, Oceania and adjacent western Pacific Rim countries. Staff scientists in the Division use a variety of approaches, including general morphology, morphometry, and molecular techniques. Biodiversity surveys and monitoring population and community structure are regular features of the staff’s fieldwork.

The Amphibian and Reptile Collection is the largest and among the most important in the world, numbering over 575,000 specimens organized alphabetically by taxonomy, and then numerically within a species. Each year about 2,000 new specimens are added to the collection and about 1,200 specimens are sent on loan to other researchers. The oldest documented specimen dates back to 1834. The collection is comprised of over 154,000 frogs, 230,000 salamanders, 370 caecilians, 800 crocodilians, 16 tuatara, 115,000 lizards, 450 amphisbaenids, 54,000 snakes, and 19,000 turtles. Of these approximately 13,000 are type specimens, with highest representation of North and Latin American taxa. The majority of specimens, 552,000, in the Amphibian and Reptile collection are wet collections – specimens stored in 70% ethanol. The division also maintains 13,400 dry collections, mostly skeletal material but also including flat skins. The glycerin-stored cleared and stained collection – specimens resulting from a process that transparently clears the specimen tissues leaving bone stained red and cartilage blue – counts about 4,200 specimens and mainly includes preparations of small specimens that would be damaged or deformed during the process of making traditional skeletal preparations. The Division has 7,850 formalin-stored specimens, primarily consisting of amphibian larvae, particularly tadpoles. The histological slide collection of 1,600 features microscope slides from Ernest Wever’s research on amphibian and reptile ears but also includes important representative slides from aging and reproductive studies. The Division has a sound archive that includes both the original and archival copies of audiotapes, primarily of frog vocalizations, as vouchers of published works and species reference. Most tapes have been digitized and transferred to CDs. Images, including print and digital photographs as well as radiographs are also included in the Division’s collections. Tissue samples, although not considered part of the permanent collections because they are typically consumed by the analysis, are routinely collected and a sizable number representing a variety of taxa are available for research and study.
Contacts: Jeremy Jacobs and Steve Gotte

Division of Birds
Research in the Division of Birds is oriented toward the evolution, biogeography, and systematics of birds. Particular interests include functional anatomy, structural adaptation, phylogeny, distribution and systematics of Neotropical birds, conservation biology of North American migrants, forensic ornithology, and paleontology and evolution of birds and of is-land avifaunas. Recent field sites include southeastern United States, Texas, California, Jamaica, Guyana, Peru, Paraguay, Uruguay, Korea, Burma and Gabon. In cooperation with the U.S. Armed Forces and Federal Aviation Administration, specialized research is currently underway in microscopic feather identification applying forensic methodologies to determine species of birds from fragmentary evidence, especially in relation to bird strikes on aircraft.

The Division of Birds maintains the third largest bird collection in the world, with approximately 600,000 specimens including many historical specimens, such as a Charles Darwin specimen that may be the only one in a North American museum – one of the few existing specimens to bear Darwin’s original field label. There are also specimens collected by Alfred Russel Wallace, William Henry Hudson, and other notables. The National Collection, known in the ornithological literature by the acronym USNM (referring to the old name of United States National Museum), has representatives of about 80% of the approximately 10,500 known species in the world’s avifauna. The first group of specimens originated from the private collection of Spencer Fullerton Baird, who collected in the Carlisle, Pennsylvania region in the early 1840′s. Baird’s collection also contained material from leading American naturalists of the early 1800′s, such as J. J. Audubon, and J. K. Townsend. The bird collection served as the repository for many of the specimens from the U. S. Exploring Expedition and of the surveys in the 1800′s to explore the western territories, railroad and telephone routes as well as international boundary surveys. Theodore Roosevelt collected birds as a young boy and also as a member of the Smithsonian African Expedition; his specimens are part of the USNM collection. A major portion of the bird collection came from the activities of the U.S. Biological Survey, which actively collected over much of North America from the 1890′s to 1930′s. The oldest known specimen in the Division was collected in Brazil in 1818. While the majority of the specimens in the Bird Division consist of study skins (about 500,000), skeletal (60,000) and anatomical (ethanol-stored: 30,000) specimens are also maintained and these represent the largest and most diverse of these types of collections in the world. The skeletal collection includes representatives of over 5,100 different taxa. The fluid-stored collection has representatives of almost 4,200 different taxa as well as specialized subsets including a collection of fluid-preserved stomach contents, brains, syringes and a small cleared and stained collection. Additional collections include egg sets (33,012), nests (4,900), and mounted skins (ca. 2,200). The collection also includes approximately 40,000 frozen tissue samples. About 1,100 specimens are added to the collections each year and 35-50 loans of specimens sent to qualified researchers, students and exhibitions. Tissues frozen in liquid nitrogen have also been preserved and are stored at the Laboratories of Analytical Biology. The bird collection includes 3,968 primary type specimens. Information and specimen data for the type specimens is available through an electronic database – the USNM Birds Type Catalog. Approximately 70% of the main collection is computerized in an internal specimen data base. The geographic coverage of the bird collection is worldwide including major holdings from North America, Central America, the West Indies, northern South America, eastern Africa, and Southeast Asia. Regions that are insufficiently represented include southern South America, western Africa, Europe, northern Asia, New Zealand and Australia and New Guinea.
Contacts: Brian Schmidt and Chris Milensky

Division of Fishes
Research in the Division of Fishes is directed primarily toward systematic revisions of species, genera, and families, and the interpretation of higher classification and biogeography. Staff research efforts are currently focused on the Caribbean and Indo-Pacific marine shore fishes, especially blennies and gobies; beloniform, scombroid, pleuronectiform fishes world-wide; larval fish studies, ontogeny and reproductive morphology; and Southeast Asia, South American and African freshwater fishes, especially atherinoid characiforms and catfishes. Osteological, myological and other studies are being conducted as a basis for understanding the phylogeny and higher classification among a broad range of taxa.

The Division of Fishes maintains the largest collection of fishes in the world with over 975,000 lots – specimens of the same species collected at the same time and place – totaling over 6 million individual specimens. The collection is arranged phylogenetically by family and then alphabetically by genus and species within each family. Over 35% of the collection has been computer catalogued and is accessible through an online searchable database. Specimens include adult fish as well as egg, larval and juvenile stages. For some taxa, especially those who progress through varied morphologies, preserved representatives of the complete series of life stages are available. The majority of specimens are stored in ethanol but the collection also includes dry skeletons (5,064) and specially prepared (cleared and stained) articulated skeletons (5,330) stored in glycerin as well as histology slides and otoliths. The collections include many rare and important fish species, including a Coelacanth, Latimeria chalumnae. About 25,000 or 75% of the over 33,000 known fish species are represented in the collection, including 19,000 lots (about 94,500 specimens) of type specimens representing 8,890 nominal species; including 6,375 primary types making this the largest such collection in the world. The fish collections include specimens from many historical expeditions including marine fishes from the Wilkes Expedition (1838) and U.S. Bureau of Fisheries trawling expeditions conducted by the Blake, Albatross, Fish Hawk and other ships in the late 1800′s and early 1900′s, the Smithsonian Biological Survey of the Canal Zone, as well as North American freshwater fishes collected during the Mississippi-Pacific Railroad and Mexican Boundary Surveys in the 1850′s and by David Starr Jordan and his students and colleagues (1860 to 1920). The collection has the world’s largest holdings of Indo-Pacific marine shore fishes and extensive coverage of Caribbean marine fishes as well as both North and South American freshwater fishes. In addition to the specimens, the collection includes illustrations and photographs (25,000 units) as well as radiographs (25,000) of fish. Contact: Jeffrey Williams

Division of Mammals
Research in the Division of Mammals is primarily concerned with systematic revisions, distribution and ecology, natural history, and functional anatomy. Staff research interests are concentrated on the mammals of Africa, Southeast Asia, and the Western Hemisphere. Studies of the systematics and ecology of marine mammals, especially whales and porpoises, of rodents, of bats, and of primates are being actively pursued.

With roughly 590,000 voucher specimens, the Division of Mammals maintains, by far, the world’s largest – nearly twice the size of the next largest – and one of the most important collections of mammals. The standard preparation is the skin and skull of which there are over 350,000 specimens. Other major holdings include 28,000 skeletons, 100,000 fluid-stored specimens, and 3,000 tanned skins. The collection includes 3,208 primary type specimens and many historically important specimens. The collections include several special subsets, among these are mammalian brains (857 specimens), male genitalia (1,700 specimens), fluid-preserved hearts (373), cleared-and-stained specimens (400) as well as karyotype slides (2,000), hair slides and bacula. Frozen tissue samples of vouchered specimens number about 4,000 with an additional 3,000 samples without vouchers.

The oldest specimens originated from the activities of the U.S. Exploring Expedition, dating from 1838-1842, and the personal collection of Spencer Fullerton Baird. A significant portion of the collection’s North American specimens resulted from the Biological Survey program, initiated by C. Hart Merriam and conducted by the U.S. Department of Agriculture, in the 1890s-1930s. The Mammal collection includes specimens from William L. Abbott who made large collections of mammals from Central and Southeast Asia. The Smithsonian African Expedition acquired many specimens from east Africa (1909-1911), some of which were collected by former President Theodore Roosevelt, and during the 1960s, large field pro-grams surveying mammals as disease vectors, such as the Smithsonian Venezuelan Project and the African Mammal Project, added more than 100,000 specimens to the collection.

Each year 1,500 specimens are loaned to qualified researchers. Data for over 546,000 specimens are electronically available through a searchable database. The taxonomic and geographic scope of the USNM mammal col-lection spans the globe, with especially strong representation from North America, Central America, northern South America, Africa, and Southeast Asia. Contacts: Darrin Lunde and Suzanne Peurach

Specialized facilities including radiographic and light photography systems (both digital and film in each case), dark-room, digital imaging and histological facilities, and sound analysis equipment are available. These are supplemented by discipline specific libraries and archives of original illustrations, maps, and sound recordings.

Field Work
Staff in the Department of Vertebrate Zoology conduct field research on all continents with particular emphasis throughout the Americas, portions of Africa and Southeast Asia and adjoining regions and across many portions of the World Ocean. In recent years traditional forms of specimen preparation have been supplemented by photographic documentation of life coloration, more encompassing anatomical preparations, and preservation of materials for molecular studies.

Education and Outreach
Graduate Programs are available in conjunction with University of Maryland and George Washington University including formal affiliations through the Robert Weintraub Program in Systematics and Evolution ( Through this program GWU faculty and graduate students work on a variety of organisms including bacteria, protists, angiosperms, cnidarians, mollusks, polychaete worms, arthropods, echinoderms, dinosaurs, fish, mammals and lizards.

Staffs in the Department of Vertebrate Zoology and affiliated agencies are also active as advisors to students throughout North America and in some countries in Central and South America and Europe. Students and researchers are welcome to conduct scientific investigations using the collections and facilities within the Department and may borrow certain materials for loan through their academic advisors and institutions.

The library holdings in Vertebrate Zoology are divided among divisional libraries with references focusing on systematics, taxonomy, anatomy and physiology, ecology and distribution, and evolution of their respective subject groups. The Birds collection has over 10,000 volumes, including approximately 100 journal subscriptions. The Fishes library has over 8,000 volumes, including 106 journal subscriptions on fish biology, and over 120,000 reprints of scientific literature on fish taxonomy and systematics. The Mammals collection contains about 4,500 volumes, including 40 journal subscriptions. The Amphibian and Reptile Library has approximately 3,500 volumes, maintains 35 journal subscriptions, and includes over 70,000 herpetological reprints making it the largest such collection in the world.

Programs and Partnerships

Genetics Program
The Genetics Program, currently housed at the National Zoological Park, uses molecular genetic methods in support of studies in systematics, population and conservation genetics, and molecular ecology. Much of the research in this lab is directly applicable to concerns of conservation biology, and relevant to endangered species and biodiversity issues. The lab has specializations in the analysis of ancient DNA, often from extinct birds and mammals; the genetics of host vector parasite interactions; and DNA typing to determine identity and relatedness of individuals, often using sub-optimal materials such as scats or hair samples. Contact: Jesus Maldonado.

Marine Mammal Program
Established in 1972, the Marine Mammal Program, which focuses on whales, dolphins, porpoises, sea cows, seals, and sea lions, is a cooperative research program whose principal goal is to extract all biological data possible from stranded and incidentally taken animals. Through a thorough examination of stranded and incidentally taken animals, valuable data is gained on many aspects of the normal life history of cetaceans. Scientists routinely collect data and specimens that relate to stomach contents, relative organ weights, parasite burden, reproductive condition and stage of physical maturity. Staff members also take external morphometrics and photographs of the external pigmentation pattern. The collection of marine mammals is the largest in the world, consisting of more than 6,400 specimens of cetaceans, 3,100 specimens of pinnipeds and 380 specimens of sirenians. Most of these are represented by osteological material although the collection also includes fluid and frozen specimens. Contact: Kristofer Helgen.

Research Staff

BALDWIN, Carole C., Research Zoologist and Curator of Fishes New World Shorefishes. B.S. (1981) James Madison University; M.S. (1986) College of Charleston; Ph.D. (1992) College of William and Mary. Research specialties: Diversity and evolution of tropical marine and deep-sea fishes; marine conservation; public communication of marine science.

BRAUN, Michael J., Research Zoologist. B.A. (1977) Cornell University; Ph.D. (1983) Louisiana State University. Research specialties: Molecular phylogenetics, molecular evolutionary genetics, avian hybridization and speciation, biogeography of Neotropical birds, conservation genetics.

DE QUEIROZ, Kevin, Research Zoologist and Curator of Amphibians and Reptiles. B.S. (1978) University of California, Los Angeles; M.S. (1985) San Di-ego State University; Ph.D. (1989) University of California, Berkeley. Research specialties: Systematics and evolutionary biology of amphibians and reptiles; principles and methods of systematic biology.

GRAVES, Gary R., Chair of Vertebrate Zoology, Research Zoologist and Curator of Birds. B.A. (1976) University of Arkansas, Little Rock; M.S. (1980) Louisiana State University; Ph.D. (1983) Florida State University. Research specialties: Evolution, biogeography, and ecology of birds and natural history of vertebrates.

HELGEN, Kristofer M., Research Zoologist and Curator of Mammals. B.A. (2001) Harvard University; Ph.D. (2007) University of Adelaide. Research specialties: Systematics, biogeography, anatomy, and conservation of terrestrial mammals worldwide, especially Australia.

JAMES, Helen F., Research Zoologist and Curator of Birds. B.A. (1977) University of Arkansas; D.Phil. (2000) Oxford University. Research specialties: Systematics, evolutionary morphology, and fossil record of birds; island biogeography and paleoecology; ecological effects of humans in island and marine ecosystems.

JOHNSON, G. David, Research Zoologist and Curator of Marine Larval Fishes. B.S. (1967) University of Texas, Austin; Ph.D. (1977) Scripps Institution of Oceanography. Research specialties: Systematics, comparative anatomy, ontogeny, phylogeny, and early life history of fishes, particularly acanthomorphs.

PARENTI, Lynne R., Research Scientist and Curator of Indo-Pacific Freshwater and Coastal Fishes. B.A. (1975) State University of New York, Stony Brook; Ph.D. (1980) City University of New York. Research specialties: Phylogenetic systematics and biogeography of tropical freshwater and coastal marine fishes, especially atherinomorphs and gobioids; comparative biogeography theory and methods; reproductive and nerve characters in fish systematics.

VARI, Richard P., Research Zoologist and Curator of Neotropical and African Freshwater Fishes. B.A. (1971) New York University; Ph.D. (1976) City University of New York. Research specialties: Systematics, evolution, and zoogeography of South American and African characiforms and some South American and Asian siluriforms.

Affiliated Research Staff

CARLETON, Michael D., Research Zoologist and Curator of Rodents. B.A. (1966) University of Massachusetts; Ph.D. (1979) University of Michigan. Research specialties: Systematics and evolution of muroid rodents.

CHESSER, Robert Terry, Adjunct Scientist, Patuxent Wildlife Research Center, U.S. Geological Survey. B.A. (1982) Georgia State University; Ph.D. (1995) Louisiana State University. Research specialties: North American birds; seasonal distribution of South American austral migrant birds; biogeography and systematics of birds; modern molecular and cladistic techniques for reconstruction of phylogeny, character evolution, and biogeographic history.

COLLETTE, Bruce B., Adjunct Scientist, Systematics Laboratory, National Marine Fisheries Service, Department of Commerce. B.S. (1956), Ph.D. (1960) Cornell University. Research specialties: Systematics, evolution, zoogeography, anatomy, and biology of marine fishes, especially Scombroidei (mackerels and tunas), Xiphioidei (bill-fishes), Beloniformes (needlefishes and halfbeaks), and Batrachoididae (toadfishes).

DOVE, Carla, Research Scientist. B.S. (1986) University of Montana; M.S. (1994), Ph.D (1998) George Mason University. Research specialties: Forensic ornithology; researches microscopic variation in downy feather structures and identifies unknown feather samples retrieved from aircraft engines, wildlife cases, prey remains, and anthropological artifacts.

EMMONS, Louise, Research Associate. B.A. (1965) Sarah Lawrence College; Ph.D. (1975) Cornell University. Research specialties: Tropical rainforest mammals, especially rodents; Neotropical forest and southern savanna mammals.

FOSTER, Mercedes S., Research Associate. B.A. (1963), M.A. (1965) University of California, Berkeley; Ph.D. (1974) University of South Florida. Research specialties: Evolution, ecology, and behavior of birds; tropical ecology; biodiversity methods; frugivorous birds, fruit nutrition, and seed dispersal.

GARDNER, Alfred L., Adjunct Scientist, Biological Resources Division, Patuxent Wildlife Research Center, U.S. Geological Survey. B.S. (1962), M.S. (1965) University of Arizona, Tucson; Ph.D. (1970) Louisiana State University, Baton Rouge. Research specialties: Systematics and nomenclature of mammals of the Western Hemisphere.

HEYER, W. Ronald, Curator Emeritus. B.A. (1963) Pacific Lutheran University; M.A. (1965), Ph.D. (1968) University of Southern California. Research specialties: Systematics, evolution, and biogeography of Neotropical amphibians.

McDIARMID, Roy W., Adjunct Scientist, Patuxent Wildlife Research Center, U.S. Geological Survey. B.A. (1961), M.S. (1966), Ph.D. (1969) University of Southern California. Research specialties: Natural history and evolution of amphibians and reptiles, especially Neo-tropical forms; morphology and evolution of amphibian eggs and larvae (tadpoles); standard methods for inventory and monitoring species; world snake diversity; bibliographic history of herpetology.

MEAD, James G., Curator of Mammals Emeritus. B.A. (1965) Yale College; M.A. (1972) University of Texas; Ph.D. (1972) University of Chicago. Research specialties: Evolution and interrelationship of cetaceans; functional anatomy, distribution, and biology of cetaceans in the western and northern Atlantic.

MUNROE, Thomas, Adjunct Scientist, Systematics Laboratory, National Marine Fisheries Service, Department of Commerce. B.A. (1973), M.S. (1976) Southeastern Massachusetts University; Ph.D. (1987) College of William and Mary, Virginia Institue of Marine Science. Research specialties: Systematics, evolution, biogeography, and biology of marine fishes, especially Pleuronectiformes.

OLSON, Storrs L., Curator of Birds Emeritus. B.A. (1966), M.S. (1968) Florida State University; Sc.D. (1972) Johns Hopkins University. Research specialties: Paleontology and systematics of birds, with emphasis on island avifaunas, evolution of seabirds, and neotropical bio-geography.

SPRINGER, Victor G., Senior Scientist Emeritus. B.A. (1948) Emory University; M.S. (1954) University of Miami; Ph.D. (1957) University of Texas. Research specialties: Systematics, zoogeography, and anatomy of tropical marine fishes.

THORINGTON, JR., Richard W., Research Zoologist and Curator of Primates and Squirrels. B.A. (1959) Princeton University; M.A. (1963), Ph.D. (1964) Harvard University. Research specialties: Systematics, ecology, and anatomy of squirrels and New World monkeys; studies of form and function; allometry and morphometrics.

WEITZMAN, Stanley H., Curator of Fishes Emeritus. B.A. (1951), M.A. (1953) University of California, Berkeley; Ph.D. (1960) Stanford University. Research specialties: Systematics, anatomy and phylogeny of South American characiform fishes.

WILSON, Don E., Curator of Mammals Emeritus. B.S. (1965) University of Arizona; M.S. (1967), Ph.D. (1970) University of New Mexico. Research specialties: Evolutionary biology of mammals, especially bats; mammal species of the world.

WOODMAN, Neal, Adjunct Scientist, Patuxent Wildlife Research Center, U.S. Geological Survey. B.A. (1980) Earlham College; M.S. (1982) University of Iowa; M.Phil. (1986), Ph.D. (1992) University of Kansas. Research specialties: Taxonomy, systematics, biogeography, morphology, and phylogenetics of mammals, especially the Soricidae; tropical mammal communities.

ZUG, George R., Curator of Amphibians and Reptiles Emeritus. B.A. (1960) Albright College; M.S. (1963) University of Florida; Ph.D. (1968) University of Michigan. Research specialties: Evolution and systematics of amphibians and reptiles, with emphasis on South Pacific species; biology and systematics of turtles.