Department of Entomology
The mission of the Department of Entomology is to describe and understand the phylogenetic and biological diversity of insects and other terrestrial arthropods through global field and laboratory research; to care for and improve the world’s largest accessible and most comprehensive terrestrial arthropod collection; and to disseminate these discoveries through scholarly and popular publication, databases of systematic and collection information, training at the graduate and post-graduate level, lectures, teaching and consulting, outreach, and through museum exhibition. The Department consists of staff from three government agencies: the Smithsonian Institution; the U.S. Department of Agriculture, Agricultural Research Service, Systematic Entomology Laboratory (SEL); and the U.S. Department of Defense, Walter Reed Army Institute of Research, Walter Reed Biosystematics Unit (WRBU). This combined community represents, by far, the greatest concentration of entomological expertise in the world.
Research in the Department of Entomology is primarily collection-based and focuses on systematics in the broadest sense, including basic taxonomy, comparative morphology, and life history of insects, as well as evolutionary and population biology, phylogenetics, biogeography, biodiversity, ecology, behavior, and molecular genetic studies. Of particular current interest are studies on the Classes Insecta and Arachnida.
The U.S. National Entomological Collection at the National Museum of Natural History (NMNH) ranks as probably the largest accessible insect collection in the world with approximately 35 million specimens including over 120,000 primary types plus secondary types. With specimens from worldwide locations, the collections are second to none in coverage for the Nearctic and Neotropical regions. Specimens from the Old World are also well represented, especially from Sri Lanka, the Philippines, China, and Papua New Guinea. Although the bulk of the collection is kept dry, various groups-such as spiders, adult aquatics insects, and insect larvae- are stored in ethanol. The collections are typically arranged by taxon; lower categories (at least genus, species) are arranged alphabetically, and for select taxa, for example Lepidoptera, within each species they are further organized by country of origin. For some groups, collections are currently being housed off-site as part of the collaborative Off-Site Enhancement Program with other institutions (see mites, Coleoptera, Diptera below). All families have been recently profiled by storage unit (drawer, jar, slide box) as to their curatorial health. There is an ever expanding image library being built for many groups, especially for the primary type specimens. The collections are supplemented by the Entomological Illustration Archive, totaling over 5,000 illustrations created to support the research publications of Department entomologists and to be available to the external scientific and public communities.
Although the U.S. National Museum (USNM) was established in 1842, the first record of an insect collection stored in the museum does not appear until 1858. In the 1860’s most of the Smithsonian’s USNM insect collection was sent to collaborating specialists with the stipulation that the material could be reclaimed at any time. In the early 1870’s the USDA was made the official repository for the Smithsonian insect collection, and then in 1881 the combined insect collection was formally transferred to the Smithsonian where it resides today.
The collections include a very large ectoparasite collection, worldwide in coverage and with important medical and veterinary entomology components; the Centers for Disease Control (CDC) collections of Anoplura and Siphonaptera; the Carriker collection of Mallophaga (containing 650 type specimens of Neotropical species); the K.C. Emerson collection of Mallophaga; the Jellison collection of ectoparasites; and projects sampling mammals in Panama, Venezuela, and Africa have produced large additions to the ectoparasite collections.
For detailed information about the Entomology collections, and individuals to contact for specific groups, visit our web site: http://www.entomology.si.edu/Collections.html.
Arachnid Collections – mites, ticks, spiders
Among the arachnid collections, the largest and most significant is the Acari (mite) Collection, currently housed at the USDA Beltsville Agricultural Research Center (BARC) in Maryland. It is the finest in existence for mites parasitizing humans, animals, and plants. The collection includes over 332,000 slides, 14,000 vials and 1,925 primary types. Some of the most important type components include: the collection of H.E. Ewing and I. M. Newell; nearly complete collection of E.W. Baker, C. E. Yunker and A.P. Jacot; important specimens of N. Banks; and type specimens representing all of the new species described by A. Fain from the Congo. The myriapod holdings rate second only to the Acarina, with special strength in New World specimens. The collection contains nearly all of the types of C.H. Bollman, R.V. Chamberlin, O.F. Cook, R.E. Crabill, R.L. Hoffman, H.F. Loomis, and J. McNeill. The Tick Collection (one million) was acquired by F.C. Bishopp and later combined with the collection of the Rocky Mountain Laboratory of the National Institutes of Health, Hamilton, MO, and contains 222 holotypes (26% of the know species in the world). Both the Tick and the Phytoseiid mite collections are housed off-site (Georgia Southern University and Florida Department of Agriculture in Gainesville, respectively) through cooperative Off-site Enhancement Program agreements. The Spider Collection counts over 200,000 specimens, mostly from the New World, and has over 300 types. Notable collectors include: N. Banks, R.V. Chamberlin, H. Exline, I. Fox, E.V. Keyserling, G. Marx, A. Petrunkevitch, and E. Simon. Contact: Floyd Shockley
Coleoptera Collections – beetles, weevils
The Coleoptera Collection, numbering about 12 million specimens including 26,000 types, includes adult and immature beetles and is the largest accessible beetle collection in the New World. The NMNH Coleoptera holdings include the historic T.L. Casey Collection, comprised of almost 117,000 specimens representing over 20,000 species, including 9,200 types. Other important material comes from the historic collections of G.H. Dieke and R. Korschefsky (Coccinellidae); F. Monros, D. Blake, I. Lopatin (Chrysomelidae); J.D. Sherman (aquatic Coleoptera); F.F. Tippman (Cerambycidae); O.L. Cartwright (Scarabaeidae and Cicindelinae); and P. Spangler (aquatic Coleoptera). SI and USDA-SEL staff have added significant, modern, well-curated specimens, including well over five million specimens collected from the canopy of Neotropical rain-forests by T. L. Erwin. The collection of beetle larvae and pupae, acquired through the efforts of A.G. Boving, is worldwide in representation and one of the largest in existence. Most Scarabaeidae are housed at the University of Nebraska, State Museum through a cooperative Off-site Enhancement Program. In 2009 the S. L. Wood Bark Beetle collection of over 80,000 specimens, including about 1,200 primary types was added making the NMNH collection the best in the world for this group. Contact: Floyd Shockley
Diptera Collections – flies, mosquitoes
The collections of Diptera rank among the most extensive in the world, with more than 8,059 drawers of pinned material, 8,538 boxes of slide-mounted specimens, 3238 jars of vials of specimens in alcohol and including about 20,500 primary types. Several large acquisitions, such as the collections of Charles P. Alexander (1.6 million crane flies), P. Arnaud (700,000), S.W. Bromley (35,000), A.L. Melander (250,000), John N. Belkin (92,000), and A.E. Pritchard (27,000), have greatly expanded coverage. Among the families particularly well rep-resented are the Asilidae, Tachinidae, Cecidomyiidae, Culicidae, Ephydroidea, and Tipulidae. The Department serves as the world center for mosquito research, hosting the Mosquitoes of Southeast Asia study and the Medical Entomology Project who have described over 100 new species of mosquitoes. The Mosquito Collection counts more than 300,000 specimens including 1,200 primary types located at the Museum Support Center (MSC) in Suitland, Maryland. The Lauxanoidea are at the California Department of Food and Agriculture under an Off-site Enhancement Program. Contact: Floyd Shockley
Hemiptera Collections – true bugs, cicadas, aphids, whiteflies, psyllids
The Hemiptera Collection (Heteroptera plus Homoptera) is the largest in the world and is located at the NMNH and at BARC (USDA). Although New World holdings predominate, the Old World holdings are rapidly expanding. The collection incorporates many important private collections including: A.C. Baker, H.G. Barber, C.K. Brian, T.D.A. Cockerell, C.J. Drake (including the H. Hacker, M.S. Pennington, C.E. Reed collections), A. Fitch, W.D. Funkhouser, F.W. Goding, H.M. Harris, F.C. Hottes, H.H. Knight, N.A. Kormilev, W.L. McAtee, T. Pergande, P.R. Uhler, R. A. Poisson, and, more recently, the J.T. Polhemus collection of aquatic and semi-aquatic Heteroptera, the J. Moldonado collection of Reduviidae, and the W. Ullrich collection. The Whitefly Collection (Aleyrodi-dae) is one of the world’s best collections, with over 32,500 microscope slide-mounts representing more than 1,100 species, and an extensive collection of dry preserved material. The collection includes more than 300 primary types. The Psyllid Collection includes both pinned (more than 20,000) and slide-mounted (more than 5,000) specimens which include more than 300 primary types. The Aphidoidea Collection is one of the largest collections of aphidoids in the world. The collection contains more than 90,000 slides representing over 2,400 species. The subset Aphid Collection contains primary type material for 747 species which includes 1380 primary type slides. The Coccoidea Collection (scale insects) consists of over 146,000 slides and has more than 280 primary types as well as a large collection of unmounted dry material containing several million specimens. Contact: Floyd Shockley
Hymenoptera Collections – ants, bees, wasps
The Hymenoptera Collection consists of about 4 million specimens including pinned specimens stored in more than 7,000 drawers, approximately 700 jars of vials of larvae and adults in alcohol, and includes over 17,000 primary types. The collection represents about 15 percent of the total entomological collections, and is especially rich in Symphyta, aculeates, and entomophagous parasitoids from worldwide locations. Outstanding holdings include the W.H. Ashmead, C.F. Baker, P.D. Hurd, Jr., K.V. Krombein, W.M. Mann, M.R. Smith, and A.W. Stelfox collections. Currently the most actively researched groups are the ants and parasitica. Contacts: Floyd Shockley
Isoptera, Orthoptera, Thysanoptera Collections – termites, grasshoppers & crickets, thrips
The Termite (Isoptera) Collection has 240,000 specimens and is the second largest in the world, including 1,150 of the known 2,000 species, and 943 types. The Grasshoppers, Katydids, Crickets (Orthoptera) have about 400,000 specimens – perhaps the 3rd largest collection in the world, about 3,000 species, and with 793 types. The Thrips (Thysanoptera) have 108,722 slides, probably 2nd largest collection in world, and 1,118 types. These collections are located at BARC in Mary-land. Contact: Floyd Shockley
Lepidoptera Collections – butterflies, moths
The Lepidoptera Collection has 2.9 million pinned and labeled specimens in 27,000+ drawers, , including about 25,000 primary types. There are about 3,000 alcohol jars with immature stages. The collection has the most complete representation of both larvae (123,000 specimens) and adults in the Western Hemisphere. Included are 131 slide cabinets containing about 100,000 microscope slides, mainly of moth genitalia. The collection is particularly rich in Nearctic and Neotropical species as well as Palearctic material for most families. The Microlepidoptera collection contains excellent coverage of Far Eastern species. Important holdings include: W. Barnes (450,000 pinned specimens), A. Blanchard (60,000), A.E. Brower (115,000), P. Dognin (50,000), D.C. Ferguson (50,000), M. Gentili (12,000), S. Issiki (16,000), E. Jackh (55,000), F. M. Jones (10,000), A. Kawabe (22,000), S. Nicolay (100,000), J. Robert (40,000), and G. B. Small (25,000). Contact: Floyd Shockley
Other important Insecta order holdings include Trichoptera, Plecoptera, Neuroptera, Mecoptera, Odonata. The Collection also includes the classes Chilopoda, Diplopoda, Arachnida, Symphyla, and Pauropoda. The Phthiraptera, Siphonaptera, Mantodea, Blattodea, Phasmatodea, Embioptera, Zoraptera, Psocoptera, and some Coleoptera families are temporarily deactived. Contact: Floyd Shockley
The Department of Entomology currently has the most modern insect collection facility in the world. Both dry and wet collections are housed in new, airtight, pest-proof, metal specimen cabinets, about half of which are on electric compactors. The collections are enhanced by specially constructed alcohol (wet collection) storage rooms and facilities for housing reprint libraries. Modern chemical storage facilities, equipment and supplies are stored in compactor systems, walk-in and reach-in freezers, critical point dryers, and ventilated sorting center all support state-of-the-art collections care. The Department has state-of-the-art digital photographic stations for use by staff, researchers and visitors. The Entomology Molecular Systematics Laboratory, a shared facility managed by WRBU at the Museum Support Center, is also available for research investigations, in addition to the facilities of the Smithsonian’s Laboratory of Analytical Biology.
Field studies are conducted in many parts of the United States, Mexico, Central and South America, the Asia-Pacific region, and, to a lesser extent, in Europe, Africa, and Australia. Museum’s entomologists currently participate in long-term biodiversity survey projects in Costa Rica (Arthropods of La Selva), Dominican Republic, Leaf Litter Arthropods of Mesoamerica, Peru, Guyana, Papua New Guinea, the Great Smoky Mountains National Park, and Kenya, among others. Past and present major projects in Sri Lanka, Ecuador, Peru, Madagascar, and Papua New Guinea have yielded millions of specimens for research. A series of canopy-fogging projects in Central and South America, initiated in 1974, has produced nearly 9 million specimens.
The Department of Entomology produces dozens of scientific publications per year, including journal articles, monographs, and books. Members of the Department traditionally serve as officers of the Entomological Society of Washington, which publishes the Proceedings of the Entomological Society of Washington and the Memoirs of the Entomological Society of Washington. Departmental staff also serve as editors of the Journal of the International Society of Hymenopterists, etc., as well as on editorial boards of other journals around the world. The Department produces a monthly newsletter, Ent News, and a quarterly Bug Dispatch that are available electronically online through the Department’s website.
Education and Outreach
The Department of Entomology has a proven history of training postdoctoral researchers as well as graduate and undergraduate students with special partnerships through the Smithsonian-USDA-University of Maryland MCSE (Maryland Center for Systematic Entomology) program. Through a variety of other cooperative arrangements staff members act both formally and informally as advisors to graduate students and occasionally teach courses at universities both locally and abroad. Department members also advise and oversee a variety of interns and volunteers. Specimens are made available to students for thesis work through loans to their academic advisors and students and researchers are welcome to visit the entomology collections and facilities to conduct their investigations on-site. The Department sponsors an entomology-focused all day family festival, open to the general public called BugFest, and participates in other Museum family festivals. Members of the Department participate in Bioblitzes locally and elsewhere in the country.
The Entomology Library contains over 23,000 volumes, including 120 journal subscriptions on insect systematics, ecology, behavior, and related areas. The collection is especially rich in the areas of taxonomy and anatomy of insects and related arthropods, especially arachnids. It is one of the best entomological libraries in North America.
Programs and Partnerships
Maryland Center for Systematic Entomology (MCSE)
Founded in 1981, the Maryland Center for Systematic Entomology (MCSE) is a consortium for research and training in the systematics of insects and allied groups. Graduate students are enrolled in the Department of Entomology, University of Maryland, with a Smithsonian or USDA-SEL scientist as co-advisor. Research focus includes tropical biology, ecology, evolutionary biology, behavior, molecular systematics, and systematic methods, in addition to the systematics and biogeography of virtually all the major groups of terrestrial arthropods. Contact: Ted Schultz
BRADY, Sean Gary, Chair of Entomology, Research Entomologist and Curator of Hymenoptera. B.A. (1990) California Polytechnic University, Pomona; M.A. (1993) California State University, Fullerton; Ph.D. (2002) University of California, Davis. Research specialties: Systematics, phylogenetics, and molecular evolution of aculeate Hymenoptera, especially bees and ants; phylogenetic methodology; social insect evolution.
CODDINGTON, Jonathan A., Lead Global Genome Initiative, Senior Research Entomologist and Curator of Arachnida and Myriapoda. B.A. (1975) Yale; M.A. (1978), Ph.D. (1984) Harvard University. Research specialties: Systematics and behavior of spiders; species richness estimation; theory and design of biological inventories. Genomics and curation of genonomic collections
DIKOW, Torsten, Research Entomologist and Curator of Diptera. M.S. (2002) Universitaet Rostock, Germany; Ph.D. (2007) Cornell University. Research specialties: Phylogeny of asiloid flies (Apioceridae, Asilidae, Mydidae) and Diptera in general using morphological and molecular evidence; revisionary taxonomy applying cybertaxonomic tools; application of specimen occurrence data to biodiversity studies; theory and methods of phylogenetic analysis.
ERWIN, Terry L., Research Entomologist and Curator of Coleoptera. B.A. (1964), M.A. (1966) San Jose State College; Ph.D. (1969) University of Alberta. Research specialties: Research specialties: Systematics, natural history, and zoogeography of world ground beetles (Carabidae); biodiversity aspects of Neotropical forest canopy insects and their allies; conservation of tropical forests.
ROBBINS, Robert K., Research Entomologist and Curator of Lepidoptera. B.A. (1969) Brown University; Ph.D. (1978) Tufts University. Re-search specialties: Systematics of Lycaenidae, evolutionary biology of butterflies, patterns of butterfly diversity.
SCHULTZ, Ted R., Research Entomologist and Curator of Hymenoptera and Chair of Entomology. B.A. (1988) University of California, Berkeley; Ph.D. (1995) Cornell University. Research specialties: Evolution and systematics of ants, especially the fungus-growing ants (tribe Attini, subfamily Myrmicinae); historical ecology and evolution of the fungus-growing behavior; theory and method of phylogenetic analysis; quantitative methods for assessing ant biodiversity.
Affiliated Research Staff
BROWN, John W., Research Associate. B.S. (1983) San Diego State University; Ph.D. (1988) University of California, Berkeley. Research specialties: Systematics and biogeography of the moth family Tortricidae (Insecta: Lepidoptera); biogeography of the peninsula of Baja California; conservation biology; and faunal inventories.
BUFFINGTON, Matthew, Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.S. (1997) University of California, Riverside; M.S. (2000) Texas A&M; Ph.D. (2005) University of California, Riverside. Research specialties: Systematics of parasitic Hymenoptera, specifically the Cynipoidea, Proctotrupoidea and Platygastroidea; molecular systematics; digital imaging techniques involving small insects.
BURNS, John M., Curator of Lepidoptera Emeritus. A.B. (1954) Johns Hopkins University; M.A. (1957), Ph.D. (1961) University of California, Berkeley. Research specialties: Systematics and evolutionary biology of butterflies (chiefly skippers, Hesperiidae) at and around the species level, with special attention to genitalia, foodplants, and DNA barcodes; biological poetry.
CHAMORRO, Lourdes. Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.S. (1998) Ohio State University; Ph.D. (2009) University of Minnesota, St. Paul. Research specialties: Systematics of Curculionoidea, specifically Dryophthorinae (palm weevils) and Eustylini (Entiminae) (broad nose weevils).
DAVIS, Donald R., Curator of Lepidoptera. B.A. (1956) University of Kansas; Ph.D. (1962) Cornell University. Research specialties: Systematics and phylogeny of the basal families of Lepidoptera including the superfamilies Tineoidea and Gracillarioidea; biology of leaf-mining and cave-dwelling moths.
FOLEY, Desmond, Adjunct Scientist, Walter Reed Biosystematics Unit. B.S. (1979) University of New England; M.S. (1984), Ph.D. (1997) University of Queensland. Research specialties: Molecular systematics, distribution modelling, biogeography, and spatial ecology of mosquitoes.
FLINT JR., Oliver S., Emeritus Research Entomologist. B.A. (1953), M.S. (1955) University of Massachusetts; Ph.D. (1960) Cornell University. Research specialties: Taxonomy and biology of the Trichoptera and Megaloptera of the New World
FURTH, David George, Research Associate. B.A. (1967) Miami University; M.S. (1969) Ohio State University; Ph.D. (1976) Cornell University. Research specialties: Systematics and biology of Leaf Beetles (Chrysomelidae), especially Flea Beetles (Alticinae).
GATES, Michael W., Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.A. (1992) Hendrix College; M.S. (1995) Oklahoma State University; Ph.D. (2000) University of California, Riverside. Research specialties: Taxonomy and systematics of Chalcidoidea (Hymenoptera), especially Eurytomidae and Eulophidae; collecting techniques, rearing and diversity of Chalcidoidea; digital imaging and image databasing.
GOLDSTEIN, Paul Z., Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.A. (1991) Harvard University; Ph.D. (1999) University of Connecticut. Research specialties: Phylogenetic systematics, life history evolution, and host plant associations of moths, especially Noctuoidea; faunal change and comparative faunistics; phylogenetic theory, molecular systematics, and species diagnostic tools.
HENRY, Thomas J., Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.A. (1971) Purdue University; M.S. (1980) Pennsylvania State University; Ph.D. (1995) University of Maryland. Research specialties: Systematics of Heteroptera (Hemiptera), especially Berytidae and Miridae.
KONSTANTINOV, Alexander S., Research Entomologist, Systematic Entomology Lab., ARS-USDA. M.A. (1981) Byelorussian State University; Ph.D. (1988) Zoological Institute, St. Petersburg. Research specialties: Systematics, comparative morphology, biogeography, and host plants relationships of leaf beetles (Chrysomelidae) with particular emphasis on flea beetles, worldwide.
KULA, Robert, Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.S. (1998) Peru State College; M.S. (2001) Texas A&M University; Ph.D. (2006) Kansas State University. Research specialties: Systematics of Ichneumonoidea, particularly parasitoids of cyclorrhaphous flies.
LINGAFELTER, Steven W., Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.S. (1989), M.S. (1991) Midwestern State University; Ph.D. (1996) University of Kansas. Research specialties: Systematics and taxonomy of Cerambycidae and Curculionidae (longhorned wood boring beetles and weevils); bionomics of Silphidae (carrion beetles).
MATHIS, Wayne N., Curator of Diptera Emeritus. B.A. (1969) Brigham Young University; Ph.D. (1976) Oregon State University. Research specialties: Systematics, biology, and zoogeography of Canacidae, Tethrnidae, and Ephydroidea, with special emphasis on Ephydridae.
McKAMEY, Stuart H., Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.S. (1985) University of California, Berkeley; M.S. (1989) North Carolina State University; Ph.D. (1994) University of Connecticut. Re-search specialties: Biosystematics of Auchenorrhyncha, principally Membracoidea (leafhoppers and treehoppers).
METZ, Mark A., Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.S. (1988) UCLA; M.S. (1995) California State University, Northridge; Ph.D. (2002) University of Illinois at Urbana-Champaign. Research specialties: Evolutionary biology, systematics, and comparative morphology of Microlepidoptera; computer and information systems and quantitative methodologies in biology.
MILLER, Gary, Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.A. (1980) Millersville State College; M.S. (1982) University of Tennessee, Knoxville; Ph.D.(1991) Auburn University. Research specialties: Systematics and taxonomy of the Aphidoidea.
MILLER, Scott E., Deputy Under Secretary for Collections and Interdisciplinary Support. B.A. (1981) University of California, Santa Barbara; Ph.D. (1986) Harvard University. Research specialties: Systematics of Lepidoptera (moths); biogeography of Pacific Basin, New Guinea, and Africa; plant-insect community ecology.
NORRBOM, Allen L., Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.A. (1980) Drexel University; M.S. (1983), Ph.D. (1985) Pennsylvania State University. Research specialties: Systematics (taxonomy, nomenclature, identification) and natural history of true fruit flies (Insecta: Diptera: Tephritidae) and related families.
RUEDA, Pollie M., Research Entomologist, Walter Reed Biosystematics Unit. B.S. (1976), M.S. (1980) University of the Philippines; Ph.D. (1984) North Carolina State University. Research specialties: Mosquito biosystematics (Culicidae, Diptera).
SOLIS, M. Alma, Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.A. (1978), M.A. (1982) University of Texas, Austin; Ph.D. (1989) University of Maryland. Research specialties: Systematics of snout moths (Pyraloidea); Pyraloidea of Neotropical areas, particularly Costa Rica.
THOMPSON, F. Christian, Research Associate. B.A. (1966), Ph.D. (1969) University of Massachusetts. Research specialties: Systematics and zoogeography of Syrphidae and related groups; zoological nomenclature; biodiversity informatics.
VANDENBERG, Natalia J., Research Associate. B.A. (1978), Ph.D. (1987) University of California, Berkeley. Research specialties: Taxonomy of larval and adult Coleoptera, especially systematics and zoogeography of Coccinellidae.
WILKERSON, Richard C., Research Associate. B.A. (1968) University of North Carolina; M.S. (1973), Ph.D. (1978) University of Florida. Research specialties: Mosquito systematics.
WOODLEY, Norman E., Research Entomologist, Systematic Entomology Lab., ARS-USDA. B.A. (1976) Washington State University; Ph.D. (1983) Harvard University. Research specialties: Taxonomy and phylogeny of Diptera, especially Brachycera, Orthorrhapa, and Oestroidea.