Research Publications


  • Rajagopalan K., DeGrandi-Hoffman G., Pruett M., Jones V.P., Coby-Harris V., Pireaud J., Curry R. Hopkins B., Northfield T.D. (2024). Warmer autumns and winters could reduce honey bee overwintering survival with potential risks for pollination services. Scientific Reports 14: 5410.


  • Zhang G., Dilday S., Kuesel R.W., Hopkins H. (2023). Phytochemicals, Probiotics, Recombinant Proteins: Enzymatic Remedies to Pesticide Poisonings in Bees. Environ. Sci. Technol. 58, 1, 54–62
  • Hopkins B.K., Long J., Naeger N.L., Sheppard W.S. (2023). Comparison of indoor (refrigerated) versus outdoor winter storage of commercial honey bee, Apis mellifera (Hymenoptera: Apidae), colonies in the Western United States. Journal of Economic Entomology. 116(4) 1063–1068.
  • Onayemi S., Olsson R., Kulhanek K., Hopkins B.K. (2023). Summer indoor queen banking as an alternative to outdoor queen banking practices. Journal of Apicultural Research. 471-477.
  • Zhang G., Olsson R., & Hopkins B.K. (2023). Strategies and techniques to mitigate the negative impacts of pesticide exposure to honey bees. Environmental Pollution. 318: 120915.


  • Reams T., & Rangel J. (2022). Understanding the enemy: A review of the genetics, behavior and chemical ecology of Varroa destructor, the parasitic mite of Apis mellifera. Journal of Insect Science. 22(1).
  • Onayemi S., Hopkins B.K., & Sheppard W.S. (2022). Elevated CO2 increases overwintering mortality of Varroa destructor (Mesostigmata: Varroidae) in honey bee (Hymenoptera: Apidae) colonies. Journal of Economic Entomology. 115(4) 1054-1058.
  • Marcelino J., Braese C., Christmon K., Evans J.D., Gilligan T., Giray T., Nearman A., Niño E.L., Rose R., Sheppard W.S., vanEngelsdorp D., & Ellis J.D. (2022). The movement of Western honey bees (Apis mellifera L.) among U.S. states and territories: history, benefits, risks, and mitigation strategies. Frontiers in Ecology and Evolution. 10:850600.
  • Bloom E. H., Oeller E.C., Olsson R.L., Brousil M.R., Schaeffer R.N., Basu S., Fu Z., & Crowder D.W. (2022). Documenting pollinators, floral hosts, and plant–pollinator interactions in U.S. Pacific Northwest agroecosystems. Ecology. 103(3): e3606.
  • Kulhanek K., Hopkins B.K. & Sheppard W.S. (2022). Comparison of oxalic acid drip and HopGuard for pre-winter Varroa destructor control in honey bee (Apis mellifera) colonies. Journal of Apicultural Research.


  • Han, J.O., Naeger N.L., Hopkins B.K., Sumerlin D., Stamets P.E., Carris L.M., & Sheppard W.S. (2021). Directed evolution of Metarhizium fungus improves its biocontrol efficacy against Varroa mites in honey bee colonies. Scientific Reports 11:10582.
  • Auth C., Hauser M., & 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. 52, 576-579.
  • Auth C. & Hopkins B.K. (2021). Nitrogen vapor immersion: An accessible alternative for honey bee (Apis mellifera L.) semen cryopreservation. Cryobiology. 100(2) 12-18.
  • Hopkins B.K. (2021). Chapter 13: Basics of Apiary Design. In Terry R. Kane et al. Honey Bee Medicine for the Veterinary Practitioner. John Wiley & Sons. 179-182.


  • Hopkins B.K., Chakrabarti P., Lucas H.M., 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.
  • Hassler E., Cazier J., 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.

2019 – 2000

  • Ostiguy N., Drummond F.A., Aronstein K., Eitzer B., Ellis J.D., Spivak M., Sheppard W.S. (2019). Honey Bee Exposure to Pesticides: A Four-Year Nationwide Study. Insects. 10(1):13.
  • Sheppard WS. & McPheron, BA. Edited by Smith D.R. (2019). Ribosomal DNA diversity in Apidae. Diversity in the Genus Apis. 89-102. CRC Press.
  • 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.
  • Stamets P.E., Naeger N.L., Evans J.D., Han J.O., Hopkins B.K., Lopez D., Moershel H.M., Nally R., Sumerlin D., Taylor A.W, Carris L.M, & Sheppard W.S. (2018) Extracts of polypore mushroom mycelia reduce viruses in honey bees.  Scientific Reports. 8, 13936.
  • Pinto M.A., Sheppard W.S., Johnson J.S., Rubink W.L., Coulson R.N., Schiff N.M., Kandemir I., & Patton J.C. (2017). Evidence that honey bee (Hymenoptera:Apidae) mitochondrial DNA haplotypes of sub-Saharan origin occurred in non-Africanized areas of the southern United States. Annals of the Entomological Society of America. 289-295.[289:HBHAOA]2.0.CO;2
  • Koch J.B., Looney C., Sheppard W.S., & Strange, J.P. (2017). Patterns of population genetic diversity and structure across bumble bee communities in the Pacific Northwest. Conservation Genetics. 507-520.
  • Gül 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.
  • Hopkins B.K., Cobey S.W., Herr C. & 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.
  • 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. Journal of Apiculture Research. 55(4) 279-283.
  • Miguel I., Garnery L., Iriondo M., Baylac M., Carmen M., Sheppard W.S, & 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) 427-440.
  • Boyle N., & Sheppard W.S. (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.S., & 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.S. (2015). Honey Bee Diversity – Races, Ecotypes and Strains. In Joe M. Graham et al. The Hive and the Honey Bee: A New Book on Beekeeping Which Continues the Tradition of Langstroth on the Hive and the Honeybee. Dadant & Sons. 53-67.
  • Lawrence T., Culbert E., Felsot A.S., Hebert V., & Sheppard, W.S. (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., Bertrand B., Legout H., Moulin S., Alburaki A., Sheppard W.S. & Garnery L.  (2013).  A fifth major genetic group among honeybees revealed in Syria. BMC Genetics. 14:117.
  • Hopkins B.K., Herr C. & 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., Smart M.D., Anelli C.M. & Sheppard W.S. (2012).  Honey bees (Apis mellifera) reared in brood combs containing high levels of pesticide residues exhibit increased susceptibility to Nosema (Microsporidian) infection.  Journal of Invertebrate Pathology. 109(3):326-329.
  • Hopkins B.K., Baker P., Sheppard W.S., & Herr C. (2011) The importance of egg-yolk in the cryopreservation of honey bee (Apis mellifera) semen. Cryobiology. 63(3):333.
  • Wu J.C., Anelli C.M. & Sheppard W.S. (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. & Sheppard W.S. (2011).  Nosema ceranae in age cohorts of the western honey bee (Apis mellifera).  Journal of Invertebrate Pathology. 109(1):148-51.
  • Cobey S.W., Sheppard W.S. & Tarpy D.R. (2011). Chapter 4: Status of Breeding Practices and Genetic Diversity in US Honey Bees in D. Sammataro & J. Yoder’s Honey Bee Colony Health. CRC Press.
  • Hopkins B.K. & Herr C. (2010). Factors affecting the successful cryopreservation of honey bee spermatozoa. Apidologie. 41 (5) 548-556.
  • Bourgeois L., Sheppard W.S., Sylvester H.A., & Rinderer T.E (2010). Genetic stock identification of Russian honey bees.  Journal of Economic Entomology.  103:917-924.
  • Delaney D.A., Meixner M.D., Schiff N.M., & Sheppard W.S. (2009).  Genetic characterization of commercial honey bee (Hymenoptera: Apidae) populations in the United States by using mitochondrial and microsatellite markers.  Annals of the Entomological Society of America. 102(4): 666-673.
  • Haddad N., Meixner M.D., Fuchs S., Migdadi H., Garnery L., Sheppard W.S. (2009). Mitochondrial DNA support for genetic reserves of Apis mellifera/syriaca in Jordan.  Journal of Apicultural Research.  48: 19-22.
  • Stucky M., Hopkins B.K., & Herr C. (2008). Cryopreservation of honey bee spermatozoa. Reproduction, Fertility and Development. 20 (1): 127-128.
  • Arias M.C., Silvestre D., Francisco F.O., Weinlich R., & Sheppard W.S. (2008).  An oligonucleotide primer set for PCR amplification of the complete honey bee mitochondrial genome. Apidologie. 39: 475-480.
  • Strange J.P., Garnery L., & Sheppard W.S. (2008). Morphological and molecular characterization of the Landes honey bee (Apis mellifera L.) ecotype for genetic conservation.  Journal of Insect Conservation.  12: 527-537.
  • Ken T., Meixner M.D., Fuchs S., Xuan Z., Shaoyu H., Kandemir I, Sheppard W.S., & Koeniger N. (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., Garnery L., Iriondo M., Garnery L., Sheppard W.S. & Estonba A. (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. Apidologie. 38: 141-155.
  • Strange J.P., Garnery L., & Sheppard W.S. (2007). Persistence of the Landes ecotype of Apis mellifera  mellifera in southwest France: confirmation of a locally adaptive annual brood cycle.  Apidologie. 38:259-267.
  • Arias M.C., Rinderer T.E., & Sheppard W.S. (2006).  Further characterization of honey bees from the Iberian Peninsula by allozyme, morphometric and mtDNA haplotype analyses.  Journal of Apicultural Research.  45:188-196.
  • Kandemir I., Kence M., Sheppard W.S., & Kence A. (2006).  Mitochondrial DNA variation in honey bee (Apis mellifera L.) populations from Turkey.  Journal of Apicultural Research.  45: 33-38.
  • Kandemir I., Meixner M.D., Ozkan A., & Sheppard W.S. (2006). Genetic characterization of honey bee (Apis mellifera cypria) populations in northern Cyprus. Apidologie. 37:547-555.
  • Kandemir I., Pinto A.M., Meixner M.D., & Sheppard W.S. (2006).  HinF-I digestion of cytochrome oxidase I region is a diagnostic test for A. m. lamarckiiGenetics and Molecular Biology.  29: 747-749.
  • Pinto M.A., Sheppard W.S., Johnson J.S., Rubink W.L., Coulson R.N., Schiff N.M., Kandemir I., & Patton J.C.  Evidence that honey bee (Hymenoptera:Apidae) mitochondrial DNA haplotypes of sub-Saharan origin occurred in non-Africanized areas of the southern United States. Annals of the Entomological Society of America. 100(2), 289-295.[289:HBHAOA]2.0.CO;2
  • Whitfield C.W., Behura S.K., Berlocher S.H., Clark A.G., Johnson J.S., Sheppard W.S., Smith D.R., Suarez A.V., Weaver D. & Tsutsui N.D. (2006). Thrice out of Africa: Ancient and resent expansions of the honey bee, Apis mellifera Science.  314:642-645.
  • Arias M.C. & Sheppard W.S. (2005). Phylogenetic relationships of honey bees (Hymenoptera:Apinae:Apini) inferred from nuclear and mitochondrial DNA sequence data.  Molecular Phylogenetics and Evolution 37(1):25-35. https:/
  • Costa M., Del Lama M.A., Melo G.A.R., & Sheppard W.S. (2003).  Molecular phylogeny of the stingless bees (Apidae, Apinae, Meliponini) inferred from mitochondrial 16S rDNA sequences.  Apidologie.  34: 73-84.
  • Diniz N.M., Soares A.E.E., & Sheppard W.S. (2003). Genetic structure of honeybee populations from southern Brazil and Uruguay.  Genetics and Molecular Biology. 26: 47-52.
  • Kamel S., Strange J.P. & Sheppard W.S. (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., Johnson J.S., Rubink W.L., Coulson R.C., Patton J.C., Sheppard W.S. (2003).  Identification of Africanized honey bee (Hymenoptera:Apidae) mitochondrial DNA: Validation of a rapid polymerase chain reaction-based assay. Annals of the Entomological Society of America.  96:679-684.[0679:IOAHBH]2.0.CO;2
  • Sheppard W.S. & Meixner M.D.  (2003). Apis mellifera pomonella, a new honey bee from the Tien Shan Mountains of Central Asia. Apidologie. 34:367-375.
  • Strange J.P., &7. Sheppard W.S. (2001).  Optimum timing of miticide applications for control of Varroa destructor (Acari: Varroidae) in Apis mellifera (Hymenoptera: Apidae) colonies in Washington State, USA. Journal of Economic Entomology. 94:1324-1331.
  • Collins A.M., Sheppard W.S. & Shimanuki H. (2000). A scientific note on the identification of honey bee semen using a mitochondrial DNA marker. Apidologie. 31:595-596.
  • Meixner M.D., Arias M.C., & Sheppard W.S. (2000). Mitochondrial DNA polymorphisms in honey bee subspecies from Kenya. Apidologie. 31: 181-190.
  • Sheppard W.S. & Smith D.R. (2000). Identification of African-derived honey bees in the Americas: A survey of methods. Annals of the Entomological Society of America. 93: 159-176.[0159:IOADBI]2.0.CO;2

Prior to 2000

  • Sheppard W.S., Rinderer T.E., Garnery L. & Shimanuki H. (1999). Further analysis of Africanized honey bee mitochondrial DNA reveals diversity of origin. Genetics and Molecular Biology. 22:73-75.
  • DeGrandi-Hoffman G., Watkins J.C., Collins A.C., Loper G.M., Martin J.H., Arias M.C., & Sheppard W.S. (1998). Queen developmental time as a factor in the Africanization of European honey bee populations (Hymenoptera: Apidae). Annals of the Entomological Society of America. 91: 52-58.
  • Sinacori A., Rinderer T.E., Lancaster V., & Sheppard W.S. (1998). A morphological and mitochondrial assessment of Apis mellifera sicula from Palermo, Italy. Apidologie. 29:481-492
  • Sheppard W.S., Arias M.C., Meixner M.D, & Grech A. (1997). Apis mellifera ruttneri, a new honey bee subspecies from Malta. Apidologie. 28:287-293.
  • Sheppard W.S. (1997). Subspecies of Apis mellifera in Honey Bee Pests, Predators and Diseases, ed. R. Morse, 3rd edition, Cornell Univ. Press.
  • Arias, M.C. & Sheppard W.S. (1996). Molecular phylogenetics of honey bee subspecies (Apis mellifera L.) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution. 5: 557-566.
  • Arias M.C., Tingek S., Kelitu A. & Sheppard W.S. (1996). Apis nuluensis (Tingek, Koeniger and Koeniger 1996) and its genetic relationships with sympatric species inferred from DNA sequences. Apidologie. 27:415-422.
  • Schiff N.M. & Sheppard W.S. (1996). Genetic differentiation in the queen breeding population of the western United States. Apidologie. 27:77-86.
  • Schiff N.M. & Sheppard W.S. (1995). Genetic analysis of commercial honey bees (Hymenoptera: Apidae) from the southern United States. Journal of Economic Entomology. 88: 1216-1220.
  • Schiff N.M., Sheppard W.S., Loper G.R., & Shimanuki H. (1994). Genetic diversity of feral honey bee (Hymenoptera: Apidae) populations in the Southern United States. Annals of the Entomological Society of America. 87:842-848.


  • Olsson R.L. (2020). Pollinator Communities Across Agricultural Gradients. A Ph.D. dissertation, Washington State University.
  • Hopkins B.K. (2014). Artificial Reproductive Techniques In Honey Bees: Sperm Cell Physiology, Semen Collection and Storage. A Ph.D. dissertation, Washington State University.

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