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Walter S. Sheppard

Professor, Graduate Faculty

Apiculturist, Thurber Memorial Chair


Areas of Interest

Population genetics & evolution of honey bees, insect introductions & mechanisms of genetic differentiation.

Dr. Sheppard also heads the Apis Molecular Systematics Laboratory.

A video report exploring the impact of pesticides on bee colonies. view


  • University of Georgia, Zoology, B.Sc., 1975
  • University of Illinois, Entomology, MSc, 1979
  • University of Illinois, Entomology, PhD, 1986
  • USDA-ARS Beneficial Insects Laboratory, Population Genetics, Postdoctoral Fellow, 1986-1988


  • 2024 – Present: Professor Emeritus, Department of Entomology, Washington State University
  • 2018 – 2024: Thurber Endowed Professor of Apiculture, Washington State University
  • 2009 – 2018 Chair, Department of Entomology, Washington State University
  • 2009-2002: Professor of Entomology, Thurber Chair, Washington State University
  • 2001-1996: Assoc. Professor of Entomology, Thurber Chair, Washington State University
  • 1996-1988: Research Entomologist, USDA Bee Research Laboratory
  • 1988-1986: Postdoctoral Fellow, USDA Beneficial Insects Laboratory
  • 1986-1980: PhD Graduate Student, Department of Entomology, UIUC
  • 1979-1977: MSc Graduate Student and Research Assistant, Departments of Entomology/Horticulture, UIUC

Teaching Activities

ENTOM 361 Honey Bee Biology

ENTOM 362 Apiculture Lab


  • Koch, J., Looney, C., Sheppard, W., & Strange, J.P. (in press). Patterns of population genetic diversity and structure across bumble bee communities in the Pacific Northwest.  Conservation Genetics.
  • Miguel, I., Garnery, L., Iriondo, M., Baylac, M., Carmen, M., Sheppard, W., & Estonba, A. 2016. Origin, evolution and conservation of the honey bee from La Palma Island (Canary Islands): molecular and morphological data.  Journal of Apicultural Research. 54(5).
  • Boyle, N., & Sheppard, W. 2016. A scientific note on seasonal levels of pesticide residues in honey bee worker tissue.  Journal of Apicultural Research.48(1): 128.
  • Koch, J., Looney, C., Sheppard, W., & Strange, J.P. 2016. Range extension of two bumble bee species (Hymenoptera:Apidae) into Olympic National Park.  Northwest Science: Official Publication of the Northwest Scientific Association. 90(2): 228-234.
  • Sheppard, W. (2015). Honey Bee Diversity – Races, Ecotypes and Strains. In Joe Graham (Eds.),  The Hive and the Honey Bee  (2015 ed., pp 53-67). Dadant and Sons.
  • Lawrence, T., Culbert, E., Felsot, A.S., Hebert, V., & Sheppard, W. (2015). Survey and Risk Assessment of Apis mellifera Exposure to Neonicotinoid Pesticides in Urban, Rural, and Agricultural Settings.  Journal of Economic Entomology, 109(2): 520-528.
  • Alburaki, M., B. Bertrand, H. Legout, S. Moulin, A. Alburaki, W.S. Sheppard and L. Garnery.  2013.  A fifth major genetic group among honeybees revealed in Syria.  BMC Genetics.  14:117.
  • Hopkins, B.K., C. Herr, and W.S. Sheppard. 2012. Sequential generations of honey bee (Apis mellifera) queens produced using cryopreserved semen. Reprod. Fert. and Dev. doi: 10.1071/RD11088
  • Wu, J.C., M. D. Smart, C.M. Anelli and W.S. Sheppard. 2012. Honey bees (Apis mellifera) reared in brood combs containing high levels of pesticide residues exhibit increased susceptibility to Nosema (Microsporidian) infection. J. Invert. Path., doi: 10.1016/j.jip.2012.01.005
  • Wu, J.C., C.M. Anelli and W.S. Sheppard. 2011. Sub-lethal effects of pesticide residues in brood comb on worker honey bee (Apis mellifera) development and longevity. Plos-One, 6(2), e14720.
  • Smart, M.D. and W.S. Sheppard. 2011. Nosema ceranae in age cohorts of the western honey bee (Apis mellifera). J. Invert. Path., doi: 10.1016/j.jip2011.09.009
  • Cobey, S. W., D. Tarpy, and W.S. Sheppard. 2011. Breeding Practices and Genetic Diversity in US Honey Bees, in Honey Bee Colony Health Issues, D. Sammataro, editor, CRC Press, Boca Raton, FL 302pp.
  • Bourgeois, L., W. S. Sheppard, H. A. Sylvester, and T. E. Rinderer. 2010. Genetic stock identification of Russian honey bees. J. Econ. Entomol. 103:917-924.
  • Delaney, D.A., Meixner, N.M. Schiff., and W. S. Sheppard. 2009. Genetic characterization of commercial honey bee (Hymenoptera: Apidae) populations in the United States by using mitochondrial and microsatellite markers. Ann. Entomol. Soc. Amer. 102: 666-673.
  • Haddad, N., M. D. Meixner, S. Fuchs, H. Migdadi, L. Garnery, W. S. Sheppard. 2009. Mitochondrial DNA support for genetic reserves of Apis mellifera syriaca in Jordan. J. Apic. Res. and Bee World. 48: 19-22.
  • Johnson, P. L., J. L. Hayes, J. Rhinehart, W. S. Sheppard, S. E. Smith. 2008. Characterization of two non-native invasive bark beetles, Scolytus schevyrewi and Scolytus multistriatus (Coleoptera: Curculionidae: Scolytinae). Can. Entomol. 140: 527-538.
  • Arias, M. C., D. Silvestre, F. O. Francisco, R. Weinlich and W. S. Sheppard. 2008. An oligonucleotide primer set for PCR amplification of the complete honey bee mitochondrial genome. Apidologie 39 : 475-480.
  • Strange, J.P., L.Garnery, and W.Sheppard. 2008. Morphological and molecular characterization of the Landes honey bee (Apis mellifera L.) ecotype for genetic conservation. J. Insect Conservation. 12: 527-537.
  • Strange, J.P, L, Garnery and W.S. Sheppard. 2007. Persistence of the Les Landes ecotype of Apis mellifera mellifera in southwest France: confirmation of a locally adaptive annual brood cycle. Apidologie. 38:259-267.
  • Ken, T, M.A., Meixner, S. Fuchs, Z. Xuan, H. Shaoyu, I. Kandemir, W. S. Sheppard, and N. Koeniger. 2007. Geographic distribution of the eastern honeybee, Apis cerana (Hymenoptera: Apidae) across ecological zones in China: morphological and molecular analyses. Systematics and Biodiversity. 5: 1-10.
  • Kandemir, I., M. D. Meixner, A. Ozkan, W.S. Sheppard. 2006. Genetic characterization of honey bee (Apis mellifera cypria) populations in northern Cyprus. Apidologie. 37:547-555.
  • Pinto, M.A., W.S. Sheppard, J.S. Johnson, W.L. Rubink, R.N. Coulson, N.M Schiff, I. Kandemir, J.C. Patton. Evidence that honey bee (Hymenoptera:Apidae) mitochondrial DNA haplotypes of sub-Saharan origin occurred in non-Africanized areas of the southern United States. In Press – Ann. Entomol. Soc. Amer.
  • Whitfield, C.W, S.K. Behura, S.H. Berlocher, A.G. Clark, J.S. Johnson, W.S. Sheppard, D.R. Smith, A.V. Suarez, D. Weaver and N. D. Tsutsui. 2006. Thrice out of Africa: Ancient and resent expansions of the honey bee, Apis mellifera. Science. 314:642-645.
  • Kandemir I, Pinto AM, Meixner MD, Sheppard WS. 2006. HinF-I digestion of Cytochrome Oxidase I region is a diagnostic test for A. m. lamarckii only? Genetics and Molecular Biology 29: 747-749.
  • Miguel, I., L. Garnery, M. Iriondo, L. Garnery, W. S. Sheppard and A. Estonba. 2007. Gene flow within the M evolutionary linage of Apis mellifera of the Pyranees, isolation by distance and the post-glacial re-colonization routes in Western Europe. 2007. Apidologie. 38: 141-155.
  • Arias, M.C, T. E. Rinderer and W. S. Sheppard. 2006. Further characterization of honey bees from the Iberian Peninsula by allozyme, morphometric and mtDNA haplotype analyses. Journal of Apicultural Research and Bee World. 45:188-196.
  • Kandemir, I., M. Kence, W. S. Sheppard, and A. Kence. 2006. Mitochondrial DNA variation in honey bee (Apis mellifera L.) populations from Turkey. J. Apicult. Res. and Bee Wld. 45: 33-38.
  • Arias, M,C. and W. S. Sheppard. 2005. Phylogenetic relationships of honey bees (Hymenoptera:Apinae:Apini) inferred from nuclear and mitochondrial DNA sequence data. Molec. Phylogenetics and Evolution 37:25-35.
  • Johnson, D. G., B. C. Carstens, W. S. Sheppard, and R. S. Zack. 2005. Phylogeny of leafhopper subgenus Errhomus (Erronus) (Hemiptera: Cicadellidae) based on mitochondrial DNA Sequences. Ann. Entomol. Soc. Amer. 98: 165-172.
  • Sheppard, W.S. and M. D. Meixner. 2003. Apis mellifera pomonella, a new honey bee from the Tien Shan Mountains of Central Asia. Apidologie 34:367-375.
  • Costa, M., M. A. Del Lama, G.A.R. Melo, and W.S. Sheppard. 2003. Molecular phylogeny of the stingless bees (Apidae, Apinae, Meliponini) inferred from mitochondrial 16S rDNA sequences. Apidologie. 34: 73-84.
  • Kamel, S., J P. Strange and W.S. Sheppard. 2003. A scientific note on hygienic behavior in Apis mellifera lamarckii and Egyptian A. m. carnica in Egypt. Apidologie. 34: 189-190.
  • Pinto, M. A., J. S. Johnson, W. L. Rubink, R. C. Coulson, J. C. Patton and W. S. Sheppard. 2003. Identification of Africanized honey bee (Hymenoptera:Apidae) mitochondrial DNA: Validation of a rapid polymerase chain reaction-based assay. Ann. Entomol. Soc. Amer. 96:679-684.
  • Silva, J. G., M. D. Meixner, B. A. McPheron, G. J. Steck and W. S. Sheppard. 2003. Recent Mediterranean fruit fly (Diptera:Tephritidae) infestations in Florida – A genetic perspective. J. Econ. Entomol. 96: 1711-1718.
  • Diniz, N. M., A. E. E. Soares, and W. S. Sheppard. 2003. Genetic structure of honeybee populations from southern Brazil and Uruguay. Genet. Mol. Biol., 26: 47-52.
  • Meixner, M.D., B.A. McPheron, J.G. Silva, G.E. Gasparich and W.S. Sheppard. 2002. The Mediterranean fruit fly in California: evidence for multiple introductions and persistent populations based on microsatellite and mitochondrial DNA variability. Molec. Ecol. 11:891-899.
  • Sheppard W.S. and T.R. Unruh. 2001. In search of a better bee. Goodfruit grower 52: 24-26.
  • Strange, J.P., and W.S. Sheppard. 2001. Treatment thresholds and timing for thecontrol of Varroa destructor in honey bee colonies in Washington State. J. Econ. Entomol. 94:1324-1333.
  • Sheppard, W.S, A. Shoukry and S. Kamel. 2001. Apis mellifera lamarckii: the honey bee of ancient Egypt in modern times. Amer. Bee J., 141: 260-263.

*Prior to 2001, W.S.S. authored 36 additional peer-reviewed publications and 7 book chapters (full list available)

2008-2011 extension publications:

W.S.S. authored 28 “Research Reviewed” extension articles in bi-monthly column in Bee Culture, a trade magazine for the beekeeper community

Synergistic Activities:

1992-2009: US Editor of the journal Apidologie, international journal of bee research (US, French and German editorial offices), WSS remains de facto editor and WSU remains as editorial lead institution

2008-2011: maintenance and operation of a Honey Bee Disease and Pest Diagnostic facility for PNW beekeepers. Current throughput is about 1500 samples per year from both commercial and hobbyist beekeepers

Graduate and Postdoctoral Advisors:

Jaycox, Elbert R., University of Illinois, Champaign, IL (retired) MS

Berlocher, Stewart H., University of Illinois, Champaign, IL (PhD)

Thesis Advisor (graduated students) and Postgraduate-Scholar Sponsor:

Arias, Maria Cristina, University of Sao Paulo, Sao Paulo, Brazil (student, postdoc, visiting scholar)

Azeredo-Espin, Annamaria, University of Campinas, Campinas, Brazil (visiting scholar)

Costa, Marco, University of Sao Carlos, Sao Carlos Brazil (PhD)

Delaney, Debbie, Washington State University, Pullman, WA (PhD)

Del Lama, Marco, University of Sao Carlos, Sao Calos, Brazil (visiting scholar)

Diniz, Nilza, University of Ribeirao Preto, Ribeirao Preto, Brazil (PhD)

Gardner, Melissa, Washington State University, Pullman WA (MS)

Garnery, Lionel, CNRS,- Gif-sur-Yvette, France (visiting scholar)

Gasparich, Gail, Towson State University, Towson, MD (postdoc)

Kahkonen, Beth, Washington State University, Pullman WA (MS)

Kamel, Soliman, Suez Canal University, Ismailia, Egypt (PhD, visiting scholar)

Kandermir, Irfan, Washington State University, Pullman WA (PhD, postdoc)

Meixner, Marina, Institut fur Bienenkunde, Oberursel, Germany (PhD, postdoc, visiting scholar)

Oskay, Devrim, Washington State University, Pullman, WA (postdoc)

Silva, Janisete, Penn State University, State College, PA (postdoc)

Soares, Espencer, University of Ribeirao Preto, Ribeirao Preto, Brazil (visiting scholar)

Strange, Jamie, Washington State University, Pullman WA (PhD)

Ben Horwath, Washington State University, Pullman WA (MS)

Sam Hapke, Washington State University, Pullman, WA (MS)

Matthew Smart, Washington State University, Pullman WA (MS)

Judy Wu, Washington State University, Pullman, WA (MS)

Thais Collet, Washington State University, Pullman, WA (PhD)

Natalie Boyle, Washington State University, Pullman WA (MS)

Brandon Hopkins, Washington State University, Pullman, WA (PhD)

Megan Taylor, Washington State University, Pullman , WA (PhD)

Dr. Sheppard outside
Dr. Steve Sheppard

Department of Entomology
Washington State University
Pullman, WA 99164-6382

Office Number: FSHN 252A

Office: 509-335-5504
Lab: 509-335-4142
Bee Building: 509-335-8598