Peter H. Adler


Peter Adler

Department of Agricultural & Environmental Sciences
130 McGinty Court
E-143 Poole Agricultural Center
Clemson University
Clemson, South Carolina 29634-0310
Phone: (864) 656-5044
FAX: (864) 656-5069 E-mail:



My research is focused on the behavior, ecology, cytogenetics, systematics, and management of insects that affect the health and welfare of humans, domestic animals, and wildlife. Current research projects include the behavioral ecology of aquatic and terrestrial arthropods, global and local biodiversity assessments of biting flies, cytogenetic analyses of black flies, taxonomic revisions and phylogenies of black flies, host-parasite interactions, economic-impact assessments of black flies, and the role of insect-borne diseases in human history.



B.S. (1976) Biology. Washington & Lee University, Lexington, VA
M.S. (1979) Zoology. Pennsylvania State University, State College, PA
Ph.D. (1983) Entomology. Pennsylvania State University, State College, PA



ADLER, P. H., J. J. Becnel & B. Moser. 2000. Molecular characterization and taxonomy of a new species of Caudosporidae (Microsporidia) from black flies (Diptera: Simuliidae), with host-derived relationships of the North American caudosporids. Journal of Invertebrate Pathology 75: 133-143.

ADLER, P. H. & B. Belqat. 2001. Cytotaxonomy of the Prosimulium hirtipes species group (Diptera: Simuliidae) in Morocco. Journal of Insect Systematics and Evolution 32: 411-418.

ADLER, P. H. & E. A. Kachvoryan. 2001. Cytogenetics of the Holarctic black fly Simulium noelleri (Diptera: Simuliidae). Canadian Journal of Zoology 79: 1972-1979.

ADLER, P. H. & J. W. McCreadie. 2002. Black flies (Simuliidae). Pp. 185-202. In G. Mullen & L. Durden (eds.). Medical and Veterinary Entomology. Academic Press, NY.

Brockhouse, C. L. & P. H. ADLER. 2002. Cytogenetics of laboratory colonies of Simulium vittatum cytospecies IS-7 (Diptera: Simuliidae). Journal of Medical Entomology 39: 293-297.

Hamada, N., J. W. McCreadie & P. H. ADLER. 2002. Species richness and spatial distributions of blackflies (Diptera: Simuliidae) in streams of Central Amazonia, Brazil. Freshwater Biology 47: 31-40.

ADLER, P. H. & W. Wills. 2003. Legacy of death: the history of arthropod-borne human diseases in South Carolina. American Entomologist 49: 216-228. [feature article].

ADLER, P. H., D. C. Currie & D. M. Wood. 2004. The Black Flies (Simuliidae) of North America. Cornell University Press, Ithaca, NY. xv + 941 pp. + 24 color plates.

ADLER, P. H. & B. Malmqvist. 2004. Predation on black flies (Diptera: Simuliidae) by the carnivorous plant Pinguicula vulgaris (Lentibulariaceae) in northern Sweden. Entomologica Fennica 15: 124-128.

Duncan, G., P. H. ADLER, K. P. Pruess & T. O. Powers. 2004. Molecular differentiation of two sibling species of the black fly Simulium vittatum (Diptera: Simuliidae) based on random amplified polymorphic DNA. Genome 47: 373-379.

Malmqvist, B., P. H. ADLER, K. Kuusela, R. W. Merritt & R. S. Wotton. 2004. Black flies in the boreal biome, key organisms in both terrestrial and aquatic environments: a review. Écoscience 11: 187-200.

Malmqvist, B., P. H. ADLER & D. Strasevicius. 2004. Testing hypotheses on egg number and size in blackflies (Diptera: Simuliidae). Journal of Vector Ecology 29: 248-256.

Malmqvist, B., D. Strasevicius, O. Hellberg, P. H. ADLER & S. Bensch. 2004. Vertebrate host specificity of wild-caught blackflies revealed by mitochondrial DNA in blood. Proceedings of the Royal Society of London B, Biology Letters 271: S152-S155.

ADLER, P. H. 2005. Black flies, the Simuliidae. Pp. 127-140. In W. C. Marquardt (ed.). Biology of Disease Vectors, 2nd edition. Elsevier Academic Press, San Diego, CA.

ADLER, P. H., D. J. Giberson & L. A. Purcell. 2005. Insular black flies (Diptera: Simuliidae) of North America: tests of colonization hypotheses. Journal of Biogeography 32: 211-220.

McCreadie, J. W., P. H. ADLER & N. Hamada. 2005. Patterns of species richness for blackflies (Diptera: Simuliidae) in the Nearctic and Neotropical regions. Ecological Entomology 30: 201-209.

McCreadie, J. W., C. E. Beard & P. H. ADLER. 2005. Context-dependent symbiosis between black flies (Diptera: Simuliidae) and trichomycete fungi (Harpellales: Legeriomycetaceae). Oikos 108: 362-370.

Nelder, M. P., P. H. ADLER & E. A. Kachvoryan. 2005. Do gut symbiotes reflect the endemism of their host black flies (Diptera: Simuliidae) in the Caucasus of Armenia? Journal of Biogeography 32: 1333-1341.

Werner, D. & P. H. ADLER. 2005. A faunistic review of the black flies (Simuliidae, Diptera) of the federal state of Sachsen-Anhalt, Germany. Abhandlungen und Berichte für Naturkunde 27: 205-245.

Finn, D. S. & P. H. ADLER. 2006. Population genetic structure of a rare high-elevation black fly, Metacnephia coloradensis, occupying lake outlet streams. Freshwater Biology 51:2240-2251.PDF

McCreadie, J. W. & P. H. ADLER. 2006. Ecoregions as predictors of lotic assemblages of blackflies (Diptera: Simuliidae). Ecography. 29: 603-613.

Nelder, M. P., C. E. Beard, P. H. ADLER, S.-K. Kim & J. W. McCreadie. 2006. Harpellales (Zygomycota: Trichomycetes) associated with black flies (Diptera: Simuliidae): world review and synthesis of their ecology and taxonomy. Fungal Diversity 22: 121-169.

ADLER, P. H., D. Roach, W. K. Reeves, J. P. Flanagan, M. E. Morrow & J. E. Toepfer. 2007. Attacks on the endangered Attwater's Prairie-Chicken (Tympanuchus cupido attwateri) by black flies (Diptera: Simuliidae) infected with an avian blood parasite. Journal of Vector Ecology 32:309-312.

Green, T. B., S. White, S. Rao, P. P. C. Mertens, P. H. ADLER & J. J. Becnel. 2007. Biological and molecular studies of a cypovirus from the black fly Simulium ubiquitum (Diptera: SImuliidae). Journal of Invertebrate Pathology 95:26-32.

ADLER, P. H. & D. C. Currie. 2008. Simuliidae. Pp. 825-845. In R. W. Merritt, K. W. Cummins & M. B. Berg (eds.). An Introduction to the Aquatic Insects of North America, 4th edition. Kendal/Hunt, Dubuque, Iowa.

Currie, D. C. & P. H. ADLER. 2008. Global diversity of black flies (Diptera: Simuliidae) in freshwater. Hydrobiologia 595:469-475.

Hamada, N., E. da Silva Pereira & P. H. ADLER. 2008. Polytene chromosomes of Simulium (Psaroniocompsa) daltanhani from Central Amazonia, Brazil. Zootaxa 1676: 57-62.

Jitklang, S., C. Kuvangkadilok, V. Baimai, H. Takaoka & P. H. ADLER. 2008. Cytogenetics and morphotaxonomy of the Simulium (Gomphostilbia) ceylonicum species group (Diptera: Simuliidae) in Thailand. Zootaxa 1917: 1-28.

McCreadie, J. W. & P. H. ADLER. 2008. Spatial distribution of rare species in lotic habitats. Insect Conservation and Diversity 1: 127-134.

Xia, Y., A. Gower, P. H. ADLER, J. VanDyk, G. Kennedy & R. Stinner. 2008. Entomology Digital Library (EntDigiL). American Entomologist 54:10-13. Link to PDF. Digital library has moved. Link to Digital Library

Ilmonen, J., P. H. ADLER, B. Malmqvist & A. Cywinska. 2009. The Simulium vernum group (Diptera: Simuliidae) in Europe: multiple character sets for assessing species status. Zoological Journal of the Linnean Society 156: 847-863.

Kim, S.-K. & P. H. ADLER. 2009. Digestive system of larval black flies (Diptera: Simuliidae): structure and phylogeny. Invertebrate Biology 128: 182-204.

Nelder, M. P., W. K. Reeves, P. H. ADLER, A. Wozniak & W. Wills. 2009. Ectoparasites and associated pathogens of free-roaming and captive animals in zoos of South Carolina. Vector-Borne and Zoonotic Diseases 9: 469-477.

Seitz, G. & P. H. ADLER. 2009. A new species of the Simulium vernum group (Diptera: Simuliidae) from the Alps of southeastern Germany. Aquatic Insects 31: 1-10.

Tangkawanit, U., C. Kuvangkadilok, V. Baimai & P. H. ADLER. 2009. Morphotaxonomy of the Simulium (Simulium) tuberosum species group (Diptera: Simuliidae) in Thailand. Zootaxa 2048: 31-46.

Tangkawanit, U., C. Kuvangkadilok, V. Baimai & P. H. ADLER. 2009. Cytosystematics of the Simulium tuberosum group (Diptera: Simuliidae) in Thailand. Zoological Journal of the Linnean Society 155: 289-315.

Tomberlin, J. K., P. H. ADLER & H. M. Myers. 2009. Development of the black soldier fly (Diptera: Stratiomyidae) in relation to temperature. Environmental Entomology 38: 930-934.

ADLER, P. H., R. A. Cheke & R. J. Post. 2010. Evolution, epidemiology, and population genetics of black flies (Diptera: Simuliidae). Journal of Molecular Epidemiology and Evolutionary Genetics of Infectious Diseases (= Infection, Genetics and Evolution) 10: 846-865.

ADLER, P. H. & D. Werner. 2010. Eugenie Ashotovna Kachvoryan (2 March 1942 - 6 December 2008): the life and career of an Armenian entomologist, with special reference to the family Simuliidae (Diptera). Studia Dipterologica 16: 73-83.

Cammack, J. A., P. H. ADLER, J. K. Tomberlin, Y. Arai & W. C. Bridges, Jr. 2010. Influence of parasitism and soil compaction on pupation of the green bottle fly, Lucilia sericata. Entomologia Experimentalis et Applicata 136: 134-141.

Huang, Y.-T., J. Phasuk, J. Chanpaisaeng & P. H. ADLER. 2010. A new species of black fly in the subgenus Simulium (Diptera: Simuliidae) from Thailand. Medical Entomology and Zoology 61: 49-58.

Nelder, M. P., D. A. Swanson, P. H. ADLER & W. L. Grogan, Jr. 2010. Biting midges of the genus Culicoides in South Carolina zoos. Journal of Insect Science 10 (55): 1-9.

Swanson, D. A. & P. H. ADLER. 2010. Vertical distribution of hematophagous Diptera in temperate forests of the southeastern U.S.A. Medical and Veterinary Entomology 24: 182-188.

ADLER, P. H., H. C. Tuten & M. P. Nelder. 2011. Arthropods of medicoveterinary importance in zoos. Annual Review of Entomology 56: 123-142.

Belqat, B., P. H. ADLER & R. W. Crosskey. 2011. Faunistic and bibliographical inventory of the blackflies (Diptera: Simuliidae) of Morocco. Zootaxa 2829: 46-58.

Henry, W., S. Thapa, P. H. ADLER, S. K. Dey & R. Varma. 2011. Cytotaxonomy of Simulium (Montisimulium) ghoomense (Diptera: Simuliidae) from the Darjeeling Hills, India. Zootaxa 2872: 49-57.

Huang, Y.-T. & P. H. ADLER. 2011. Chromosomal relationships of Simulium suzukii, an Old World member of the Simulium tuberosum species group (Diptera: Simuliidae). Medical Entomology and Zoology 62: 23-30.

McCreadie, J. W., P. H. ADLER & C. E. Beard. 2011. Ecology of symbiotes of larval black flies (Diptera: Simuliidae): distribution, diversity, and scale. Environmental Entomology 40: 289-302.

Porter, T. M., W. Martin, T. Y. James, J. E. Longcore, F. H. Gleason, P. H. ADLER, P. M. Letcher & R. Vilgalys. 2011. Molecular phylogeny of the Blastocladiomycota (Fungi) based on nuclear ribosomal data. Fungal Biology 115: 381-392.

Pramual, P., K. Wongpakam & P. H. ADLER. 2011. Cryptic biodiversity and phylogenetic relationships revealed by DNA barcoding of Oriental black flies in the subgenus Gomphostilbia (Diptera: Simuliidae). Genome 54: 1-9.

Tang, X., P. H. ADLER, H. Vogel & L. Ping. 2012. Gender-specific bacterial composition of black flies (Diptera: Simuliidae). FEMS Microbiology Ecology 80:659-670. Link to document

King, R. S. & P. H. ADLER. 2012. Development and evaluation of methods to measure populations of black flies (Diptera: Simuliidae) at nests of the endangered whooping crane (Grus americana). Journal of Vector Ecology 37:298-306.

McCreadie, J. W. & P. H. ADLER. 2012. Variation in larval fitness of a black fly species (Diptera: Simuliidae) over heterogeneous habitats. Aquatic Insects 34: 143-150.

Swanson, D. A., P. H. ADLER & B. Malmqvist. 2012. Spatial stratification of host-seeking Diptera in boreal forests of northern Europe. Medical and Veterinary Entomology 26: 56-62.

Monaenkova, D., M. S. Lehnert, T. Andrukh, C. E. Beard, B. Rubin, A. Tokarev, W.-K. Lee, P. H. ADLER & K. G. Kornev. 2012. Butterfly proboscis: combining a drinking straw with a nanosponge facilitated diversification of feeding habits. Journal of the Royal Society Interface 9: 720-726. [Featured in Science News].

ADLER, P. H., Y.-T. Huang & H. Takaoka. 2012. Nearctic-Palearctic relationships of black flies (Diptera: Simuliidae): chromosomal and morphological evidence for the Prosimulium magnum species group in Japan. Journal of Natural History 46:1467-1475.

Tuten, H. C., W. C. Bridges, K. S. Paul & P. H. ADLER. 2012. Blood-feeding ecology of mosquitoes in zoos. Medical and Veterinary Entomology 26: 407-416.

Gray, E. W., R. D. Wyatt, P. H. ADLER, J. Smink, J. E. Cox & R. Noblet. 2012. The lack of effect of low temperature and high turbidity on operational Bacillus thuringiensis subsp. israelensis activity against larval black flies (Diptera: Simuliidae). Journal of the American Mosquito Control Association 28: 134-136.

Seitz, G., H. Zwick & P. H. ADLER. 2012. Description of a new species, Simulium (Rubzovia) joanae (Diptera: Simuliidae), and its chromosomes, with new information on the blackfly fauna of Madeira. Lauterbornia 74: 1-28.

Pramual, P., C. Kuvangkadilok, S. Jitklang, U. Tangkawanit & P. H. ADLER. 2012. Geographical versus ecological isolation of closely related black flies (Diptera: Simuliidae) inferred from phylogeny, geography, and ecology: implications for speciation in black flies (Diptera: Simuliidae). Organisms, Diversity and Evolution 12: 183-195.

Tuten, H. C., W. C. Bridges & P. H. ADLER. 2012. Comparative morphology of the pyloric armature of adult mosquitoes (Diptera: Culicidae). Arthropod Structure and Development 41: 475-481.

McCreadie, J. W. & P. H. ADLER. 2012. The roles of abiotic factors, dispersal, and species interactions in structuring stream assemblages of black flies (Diptera: Simuliidae). Aquatic Biosystems 8: 14 [11 pp.] [Editors’ Pick].

Nelder, M. P., W. K. Reeves, G. R. Curler, B. R. Ellis & P. H. ADLER. 2012 [2011]. Filth flies (Diptera) in zoos of South Carolina, USA. Studia Dipterologica 18: 11-19.

Moncada, L. A., S. Mantilla, I. Lotta, N. Matta & P. H. ADLER. 2013. Simulium cormonsi Wygodzinsky (Diptera: Simuliidae) in Colombia: first record, egg and larval descriptions, and bionomics. Proceedings of the Entomological Society of Washington 115: 103-106.

Tokarev, A., B. Kaufman, Y. Gu, T. Andrukh, P. H. ADLER & K. G. Kornev. 2012. Probing viscosity of nanoliter droplets of butterfly saliva by magnetic rotational spectroscopy. Applied Physics Letters 102: 4 pp. doi: 10.1063/1.4788927.

Lehnert, M.S., D. Monaenkova, T. Andrukh, C. E. Beard, P. H. ADLER & K. G. Kornev. 2013. Hydrophobic-hydrophilic dichotomy of the butterfly proboscis. Journal of the Royal Society Interface. [Featured in Science News].

Khazeni, A., P. H. ADLER, Z. Telmadareiiy, M. A. Oshaghi, H. Vatandoost, S. M. Abtahi & A. Lofti. 2013. The black flies (Diptera: Simuliidae) of Iran. Zootaxa 3694: 67-74.

Mantilla, J. S., L. I. Moncada, N. E. Matta & P. H. ADLER. 2013. Two new species of black flies (Diptera: Simuliidae) from the High Andes of Colombia. Zootaxa 3700: 423-434.

ADLER, P. H., Y.-T. Huang, W. K. Reeves, S. K. Kim, Y. Otsuka & H. Takaoka. 2013. Macrogenomic evidence for the origin of the black fly Simulium suzukii (Diptera: Simuliidae) on Okinawa Island, Japan. PLOS ONE 8 (8): e70765. doi:10.1371/journal.pone.0070765