• Han, J. et al. (2021) Directed evolution of Metarhizium fungus improves its biocontrol efficacy against Varroa mites in honey bee colonies. Scientific Reports 11:10582
  • Auth, C., Hauser, Hopkins, B.K. 2021. A scientific note on a black soldier fly (Stratiomydae: Hermetia illucens) infestation within a western honey bee (Apis mellifera) colony. Apidologie.


  • Hopkins, B.K., Chakrabarti , P., Lucas, H., Sagili, R., Sheppard W.S. (2020) Impacts of different winter storage conditions on the physiology of diutinus honey bees (Apis mellifera L.) Journal of Economic Entomology. 114(1) 409-414 1093/jee/toaa302
  • Sheppard, WS., McPheron, BA. Ribosomal DNA diversity in Apidae. Diversity in the genus Apis, 89-102
  • Cazier, J., Hassler, E., Hopkins, B. K., Wilkes, J. T., Smith, K., Runzel, M. A Century of Discovery: Mining 100 Years of Honey Bee Research. Journal of Apicultural Research. 60 (1) 3-12


  • Rajamohan, A., Danka, R., Hopkins, B. K., Rinehart, J. (2019). A non-activating diluent to prolong in vitro viability of Apis mellifera spermatozoa: Effects on cryopreservation and on egg fertilization. Cryobiology. (impact factor- 2.14)


  • Stamets, P., Naeger, N., Evans, J., Han, J., Hopkins, B.K, Lopez, D, Moershel, H., Nally, R., Sumerlin, D., Taylor, A., Carris, L., Sheppard, W. (2018) Extracts of polypore mushroom mycelia reduce viruses in honey bees.  Scientific Reports. 8 article# 13936 (impact factor – 4.5)


  • Pinto, M.A., Sheppard, W S, 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.
  • Koch, J., Looney, C., Sheppard, W., & Strange, J.P. (2017). Patterns of population genetic diversity and structure across bumble bee communities in the Pacific Northwest. Conservation Genetics.
  • Gul, A., Onal, A.G., Sahinler, N., Hopkins, B.K., Sheppard, W.S. (2017) Effects of frozen-thawed honey bee semen supplemented with various plasma and diluents on fertilization rate. Theriogenology. 101 109-113 (impact factor 2.29)
  • Hopkins, B.K., Herr, C. and Sheppard, W.S. 2017 Gel-coated tubes extend above-freezing storage of honey bee (Apis mellifera) semen to 439 days with production of fertilized offspring. Reproduction, Fertility and Development. 29(10) 1944-1949 (impact factor -1.7)


  • Emamverdi, M., Zhandi, M., Hopkins, B.K. (2016) Preservation of honey bee spermatozoa using egg yolk and soybean lecithin-based semen extenders and a modified cryopreservation protocol. 55(4) Journal of Apiculture Research 279-283
  • 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.

Prior to 2015

  • 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.
  • Hopkins, B.K. 2014 Artificial Reproductive Techniques In Honey Bees: Sperm Cell Physiology, Semen Collection And Storage. ProQuest. Dissertation Pub# 3628836
  • Lawrence, T., W.S. Sheppard. 2013. Neonicotinoid Pesticides and Honey Bees (PDF). WSU Extension factsheet 122E
  • 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., Herr, C. and Sheppard, W.S. 2012 Sequential generation of honey bee (Apis mellifera) queens produced using cryopreserved semen. Reproduction, fertility and Development 24(8) 1079-83
  • 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
  • Hopkins, B.K., Baker, P., Sheppard, W.S., and Herr, C. 2011 Value of egg-yolk in the cryopreservation of honey bee (Apis mellifera) semen. Cryobiology1016/j.cryobiol.2011.09.102
  • 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.
  • Hopkins, B.K. and Herr, C. (2010) Factors affecting the successful cryopreservation of honey bee semen. Apidologie 41 (5) 548-556
  • 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.
  • Hopkins B.K. and Herr, C. (2008) Cryopreservation of frog (Rana pipiens) sperm cells collected by non-lethal methods. Reproduction, Fertility and Development 20 (1): 120
  • Stucky, M., Hopkins, B.K., and Herr, C. (2008) Cryopreservation of honey bee spermatozoa. Reproduction, Fertility and Development 20 (1): 127-128
  • 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.
  • 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.
  • 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.
  • 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.
  • 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.  Journal of Apicultural Research and Bee World.  45: 33-38.
  • 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.
  • 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.
  • 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.
  • 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.
  • 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.
  • 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.
  • 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.
  • 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.
  • 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.
  • 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. 2002. Diversity of Africanized honey bees in the United States and the utility of mitochondrial DNA origins. In Africanized honey bees and bee mites II. ed. by E. Erickson. Westview Press pp 60-64.
  • Sheppard W.S. and T.R. Unruh.  2001.  In search of a better bee.  Goodfruit grower  52: 24-26.
  • 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.
  • Sheppard, W.S and  S. Kamel.  2001.  Apis mellifera lamarckii: the honey bee of ancient Egypt in modern times.  Amer. Bee J.  141:260-263
  • 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.
  • Collins, A.M., W.S. Sheppard, W.S. and H. Shimanuki. 2000. A scientific note on the identification of honey bee semen using a mitochondrial DNA marker. Apidologie 31:595-596
  • Loper, G.R., J. Fewell, E. Erickson, W.S. Sheppard. 2000. Impact of mites on, and the introgression of Africanized bees into a feral population of honey bees. Hoopingarner Roger and Conner, Lawrence J. (editors) Cheshire: Wicwas Press, UC pp. 47-51.
  • Meixner, M. D., M. C. Arias, W. S. Sheppard. 2000. Mitochondrial DNA polymorphisms in honey bee subspecies from Kenya. Apidologie. 31: 181-190.
  • Sheppard, W.S. and D.R. Smith. 2000. Identification of African-derived honey bees in the Americas: A survey of methods. Ann Entomol. Soc. Amer. 93: 159-176.
  • Sheppard, W.S., T.E. Rinderer, L. Garnery and H. Shimanuki. 1999. Further analysis of Africanized honey bee mitochondrial DNA reveals diversity of origin. Genet. and Mol. Biol. 22:73-75.
  • DeGrandi-Hoffman, G., J.C. Watkins, A. C. Collins, G. M. Loper, J. H. Martin, M. C. Arias and W. S. Sheppard. 1998. Queen developmental time as a factor in the Africanization of European honey bee populations (Hymenoptera: Apidae). Ann. Entomol. Soc. Amer. 91: 52-58.
  • Sinacori, A., T.E. Rinderer, V. Lancaster and W.S. Sheppard. 1998 A morphological and mitochondrial assessment of Apis mellifera sicula from Palermo, Italy. Apidologie 29:481-492
  • Gasparich, G.E., J.G. Silva, H. Y. Han, B.A. McPheron, G.J. Steck and W.S.Sheppard. 1997. Population genetic structure of Mediterranean fruit fly (Diptera:Tephritidae) and implications for worldwide colonization patterns. Ann. Entomol. Soc. Amer. 90:790-797.
  • Sheppard, W.S., M.C. Arias, M.D. Meixner and A. Grech. 1997. Apis mellifera ruttneri, a new honey bee subspecies from Malta. Apidologie 28:287-293.
  • Sheppard, W. S. Subspecies of Apis mellifera. 1997. in Honey Bee Pests, Predators and Diseases, ed. R. Morse, 3rd edition, Cornell Univ. Press, Arias, M.C. and W.S. Sheppard. 1996. Molecular phylogenetics of honey bee subspecies (Apis mellifera L.) inferred from mitochondrial DNA sequences. Mol. Phylogen. Evol. 5: 557-566.
  • Arias, M.C., S. Tingek, A. Kelitu and W.S. Sheppard. 1996. Apis nuluensis (Tingek, Koeniger and Koeniger 1996) and its genetic relationships with sympatric species inferred from DNA sequences. Apidologie 27:415-422.
  • Azeredo-Espin, A.M.L., Schroder, R.F.W., Roderick, G.K. and W.S. Sheppard. 1996. Intraspecific mitochondrial DNA variation in the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). Biochem. Genet. 34:253-268
  • Schiff, N.M. and W.S. Sheppard. 1996. Genetic differentiation in the queen breeding population of the western United States. Apidologie 27:77-86. Schiff, N.M. and W.S. Sheppard. 1995.
  • Genetic analysis of commercial honey bees (Hymenoptera: Apidae) from the southern United States. J. Econ. Entomol. 88: 1216-1220.
  • Schiff, N.M., Sheppard, W.S., Loper, G.R., and H. Shimanuki. 1994. Genetic diversity of feral honey bee (Hymenoptera: Apidae) populations in the Southern United States. Ann. Entomol. Soc. Amer. 87:842-848.

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