Research Publications

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Galbraith, D.A., Soojin, V.Y. and Grozinger, C.M. "Evaluation of Possible Proximate Mechanisms Underlying the Kinship Theory of Intragenomic Conflict in Social Insects." Integrative and Comparative Biology, 56(6): 1206-1214. (2016) (link)

Kammerer, M.A., Biddinger, D.J., Rajotte, E.G. and Mortensen, D.A., “Local Plant Diversity Across Multiple Habitats Supports a Diverse Wild Bee Community in Pennsylvania Apple Orchards.” Environmental Entomology. 45(1): 32-38 (2016). (link)

Padilla, M., Amsalem, E., Altman, N., Hefetz, A. and Grozinger, C.M. “Chemical communication is not sufficient to explain reproductive inhibition in the bumblebee Bombus impatiens.” Open Science. 3(10): 160576 (2016). (link)

Joshi, N.K., Otieno, M., Rajotte, E.G., Fleischer, S.J. and Biddinger, D.J. “Proximity to Woodland and Landscape Structure Drives Pollinator Visitation in Apple Orchard Ecosystem.” Front. Ecol. Evol. 4: 38  (2016). (link)

Kammerer, M.A., Biddinger, D.J., Joshi, N.K., Rajotte, E.G. and Mortensen, D.A. “Modeling local spatial patterns of wild bee diversity in Pennsylvania apple orchards.” Landscape Ecology. 31(10): 2459-2469 (2016). (link)

Bohnenblust, E.W., Vaudo, A.D., Egan, J.F., Mortensen, D.A. and Tooker, J.F. “Effects of the herbicide dicamba on nontarget plants and pollinator visitation.” Environmental Toxicology and Chemistry. 35(1): 144-151 (2016). (link)

Li, W., Evans, J.D., Huang, Q., Rodríguez-García, C., Liu, J., Hamilton, M., Grozinger, C.M., Webster, T.C., Su, S. and Chen, Y.P. “Silencing the Honey Bee (Apis mellifera) Naked Cuticle Gene (nkd) Improves Host Immune Function and Reduces Nosema ceranae Infections.” Applied and Environmental Microbiology. 82(22): 6779-6787 (2016). (link)

Holt, H.L. and Grozinger, C.M. “Approaches and challenges to managing Nosema (Microspora: Nosematidae) parasites in honey bee (Hymenoptera: Apidae) colonies.” Journal of Economic Entomology. 109(4): 1487-1503 (2016). (link)

Vaudo, A.D., Patch, H.M., Mortensen, D.A., Tooker, J.F. and Grozinger, C.M. “Macronutrient ratios in pollen shape bumble bee (Bombus impatiens) foraging strategies and floral preferences.” Proceedings of the National Academy of Sciences. 113(28): 4035-4042 (2016). (link)

Grozinger, C. “Cooperation and conflict in social insect societies: from genes to pheromones.” Integrative and Comparative Biology. 56: 81 (2016).

Rittschof, C.C., Grozinger, C.M. and Robinson, G.E. “Social cues and diet restriction may act through similar mechanisms to affect aggression in honey bees (Apis mellifera).” Integrative and Comparative Biology. 56: 184 (2016).

Galbraith, D.A., Kocher, S.D., Glenn, T., Albert, I., Hunt, G.J., Strassmann, J.E., Queller, D.C. and Grozinger, C.M. “Testing the kinship theory of intragenomic conflict in honey bees (Apis mellifera).” Proceedings of the National Academy of Sciences. 113(4): 1020-1025 (2016). (link)

Vaudo, A.D., Stabler, D., Patch, H.M., Tooker, J.F., Grozinger, C.M. and Wright, G.A. “Bumble bees regulate their intake of the essential protein and lipid pollen macronutrients.” Journal of Experimental Biology. (2016). (link)

Manfredini, F., Shoemaker, D. and Grozinger, C.M. “Dynamic changes in host–virus interactions associated with colony founding and social environment in fire ant queens (Solenopsis invicta).” Ecology and Evolution. 6(1): 233-244 (2016). (link)

Brasero, N., Martinet, B., Lecocq, T., Lhomme, P., Biella, P., Valterova, I., Urbanova, K., Cornalba, M., Hines, H. and Rasmont, P. “The cephalic labial gland secretions of two socially parasitic bumblebees Bombus hyperboreus (alpinobombus) and Bombus inexspectatus (thoracobombus) question their inquiline strategy.” Insect Science. (2016). (link)

Nadeau, N.J., Pardo-Diaz, C., Whibley, A., Supple, M.A., Saenko, S.V., Wallbank, R.W., Wu, G.C., Maroja, L., Ferguson, L., Hanly, J.J. and Hines, H. “The gene cortex controls mimicry and crypsis in butterflies and moths.” Nature. 534(7605): 106-110 (2016). (link)

Youngsteadt, E., Moylett, H., López-Uribe, M.M. and Hamblin, A. “TB Mitchell: The Man Behind The Bees of the Eastern United States.” American Entomologist. 62(3): 157-162 (2016). (link)

López-Uribe, M.M., Sconiers, W.B., Frank, S.D., Dunn, R.R. and Tarpy, D.R. “Reduced cellular immune response in social insect lineages.” Biology Letters. 12(3): 20150984 (2016). (link)

Traynor, K.S., Pettis, J.S., Tarpy, D.R., Mullin, C.A., Frazier, J.L. and Frazier, M. “In-hive Pesticide Exposome: Assessing risks to migratory honey bees from in-hive pesticide contamination in the Eastern United States.” Scientific Reports. 6:33207 (2016). (link)

Mullin, C.A., Fine, J.D., Reynolds, R.D. and Frazier, M.T. “Toxicological risks of agrochemical spray adjuvants: organosilicone surfactants may not be safe.” Frontiers in Public Health. 4:92 (2016). (link)

Russo, L. and Shea, K. “Deliberately increased network connectance in a plant-pollinator community experiment.” Journal of Complex Networks. (2016) (link)

Russo, L., Nichol, C. and Shea, K. “Pollinator floral provisioning by a plant invader: quantifying beneficial effects of detrimental species.” Diversity and Distributions. 22(2): 189-198 (2016). (link)

Feazel-Orr, H.K., Catalfamo, K.M., Brewster, C.C., Fell, R.D., Anderson, T.D. and Traver, B.E. “Effects of Pesticide Treatments on Nutrient Levels in Worker Honey Bees (Apis mellifera)." Insects. 7(1): 8 (2016). (link)

Russo, L. “Positive and Negative impacts of Non-Native Bee Species around the World.” Insects. 7 (4): 69 (2016). (link)


Amsalem, E., Grozinger, C.M., Padilla, M. and Hefetz, A. “Chapter Two-The Physiological and Genomic Bases of Bumble Bee Social Behaviour.” Advances in Insect Physiology: 48: 37-93 (2015). (link)

Amsalem, E., Grozinger, C.M., Padilla, M. and Hefetz, A. “Bumble Bee Sociobiology: The Physiological and Genomic Bases of Bumble Bee Social Behaviour.” Advances in Insect Physiology. 48: 37-93 (2015). (link)

Amsalem, E., Orlova, M. and Grozinger, C.M. “A conserved class of queen pheromones? Re-evaluating the evidence in bumblebees (Bombus impatiens).” In Proc. R. Soc. B. 282 (1817): 20151800 (2015). (link)

Amsalem, E., Galbraith, D.A., Cnaani, J., Teal, P.E. and Grozinger, C.M. “Conservation and modification of genetic and physiological toolkits underpinning diapause in bumble bee queens.” Molecular Ecology. 24(22): 5596-5615 (2015). (link)

Biddinger, D.J. and Rajotte, E.G. “Integrated pest and pollinator management—adding a new dimension to an accepted paradigm.” Current Opinion in Insect Science. 10: 204-209 (2015). (PDF)

Joshi, N.K., Leslie, T., Rajotte, E.G., Kammerer, M.A., Otieno, M. and Biddinger, D.J. “Comparative trapping efficiency to characterize bee abundance, diversity, and community composition in apple orchards.” Annals of the Entomological Society of America. 108(5): 785-799 (2015). (link)

Vaudo, A.D., Tooker, J.F., Grozinger, C.M. and Patch, H.M. “Bee nutrition and floral resource restoration.” Current Opinion in Insect Science. 10: 133-141 (2015). (PDF)

Galbraith, D.A., Wang, Y., Amdam, G.V., Page, R.E. and Grozinger, C.M. “Reproductive physiology mediates honey bee (Apis mellifera) worker responses to social cues.” Behavioral Ecology and Sociobiology. 69(9): 1511-1518 (2015). (link)

Rittschof, C.C., Grozinger, C.M. and Robinson, G.E. “The energetic basis of behavior: bridging behavioral ecology and neuroscience.” Current Opinion in Behavioral Sciences. 6: 19-27 (2015). (link)

Richards, J., Carr-Markell, M., Hefetz, A., Grozinger, C.M. and Mattila, H.R. “Queen-produced volatiles change dynamically during reproductive swarming and are associated with changes in honey bee (Apis mellifera) worker behavior.” Apidologie. 46(6): 679-690 (2015). (link)

Döke, M.A., Frazier, M. and Grozinger, C.M. “Overwintering honey bees: biology and management.” Current Opinion in Insect Science. 10: 185-193 (2015). (PDF)

Grozinger, C.M. and Robinson, G.E. “The power and promise of applying genomics to honey bee health.” Current Opinion in insect Science. 10: 124-132 (2015). (PDF)

Kocher, S.D., Tsuruda, J.M., Gibson, J.D., Emore, C.M., Arechavaleta-Velasco, M.E., Queller, D.C., Strassmann, J.E., Grozinger, C.M., Gribskov, M.R., San Miguel, P. and Westerman, R. “A search for parent-of-origin effects on honey bee gene expression.” G3: Genes| Genomes| Genetics. 5(8): 1657-1662 (2015). (link)

Fuller, Z.L., Niño, E.L., Patch, H.M., Bedoya-Reina, O.C., Baumgarten, T., Muli, E., Mumoki, F., Ratan, A., McGraw, J., Frazier, M. and Masiga, D. “Genome-wide analysis of signatures of selection in populations of African honey bees (Apis mellifera) using new web-based tools.” BMC genomics. 16(518) (2015). (link)

Villar, G., Baker, T.C., Patch, H.M. and Grozinger, C.M., “Neurophysiological mechanisms underlying sex-and maturation-related variation in pheromone responses in honey bees (Apis mellifera).” Journal of Comparative Physiology A. 201(7): 731-739 (2015). (link)

Rittschof, C.C., Grozinger, C.M. and Robinson, G.E. “Social context during pre-adult stages influences aggression in adult honey bees.” Integrative and Comparative Biology. 55: 153 (2015).

Galbraith, D.A., Yang, X., Niño, E.L., Yi, S. and Grozinger, C. “Parallel epigenomic and transcriptomic responses to viral infection in honey bees (Apis mellifera).” PLoS Pathogens. 11(3): e1004713. (link)

Cappa, F., Beani, L., Cervo, R., Grozinger, C. and Manfredini, F. “Testing male immunocompetence in two hymenopterans with different levels of social organization: ‘live hard, die young?’” Biological Journal of the Linnean Society. 114(2): 274-278 (2015). (link)

Rittschof, C.C., Coombs, C.B., Frazier, M., Grozinger, C.M. and Robinson, G.E. “Early-life experience affects honey bee aggression and resilience to immune challenge.” Scientific Reports. 5: 15572 (2015). (link)

Grozinger, C.M. and Evans, J.D. “Editorial Overview: Social insects: From the lab to the landscape-translational approaches to pollinator health.” Current Opinion in Insect Science. (10): vii-ix (2015). (PDF)

Supple, M.A., Papa, R., Hines, H.M., McMillan, W.O. and Counterman, B.A. “Divergence with gene flow across a speciation continuum of Heliconius butterflies.” BMC Evolutionary Biology. 15(1): 204 (2015). (link)

Condamine, F.L. and Hines, H.M. “Historical species losses in bumblebee evolution.” Biology Letters. 11(3): 20141049 (2015). (link)

Mullin, C.A. “Effects of ‘inactive’ ingredients on bees.” Current Opinion in Insect Science. 10: 194-200 (2015). (link)

Frazier, M.T., Mullin, C.A., Frazier, J.L., Ashcraft, S.A., Leslie, T.W., Mussen, E.C. and Drummond, F.A. “Assessing honey bee (Hymenoptera: Apidae) foraging populations and the potential impact of pesticides on eight US crops.” Journal of Economic Entomology. 108(5): 2141-2152 (2015). (link)

Mullin, C.A., Chen, J., Fine, J.D., Frazier, M.T. and Frazier, J.L. “The formulation makes the honey bee poison.” Pesticide Biochemistry and Physiology. 120: 27-35 (2015). (link)

Chen, J. and Mullin, C.A. “Characterization of trisiloxane surfactants from agrochemical adjuvants and pollinator-related matrices using liquid chromatography coupled to mass spectrometry.” Journal of Agricultural and Food Chemistry. 63(21): 5120-5125 (2015). (link)

Campbell, C., Yang, S., Albert, R. and Shea, K. “Plant–pollinator community network response to species invasion depends on both invader and community characteristics.” Oikos. 124(4): 406-413 (2015). (link)

Ellis, K.E. and Barbercheck, M.E. “Management of Overwintering Cover Crops Influences Floral Resources and Visitation by Native Bees.” Environmental Entomology. 5: 999-1010 (2015). (link)

Rangel, J., Baum, K., Rubink, W.L., Coulson, R.N., Johnston, J.S. and Traver, B.E. “Prevalence of Nosema species in a feral honey bee population: a 20-year survey.” Apidologie. 47: 561 (2015). (link)

Traver, B.E. and Fell, R.D. “A scientific note: Survey for Nosema spp. in preserved Apis spp.” Apidologie. 46(2): 194-196 (2015). (link)


Shpigler, H., Amsalem, E., Huang, Z.Y., Cohen, M., Siegel, A.J., Hefetz, A. and Bloch, G. “Gonadotropic and physiological functions of juvenile hormone in bumblebee (Bombus terrestris) workers.” PLoS One. 9(6):e100650 (2014). (link)

Amsalem, E., Kiefer, J., Schulz, S. and Hefetz, A.”The effect of caste and reproductive state on the chemistry of the cephalic labial glands secretion of Bombus terrestris.” Journal of Chemical Ecology. 40(8): 900-912 (2014). (link)

LeBoeuf, A.C. and Grozinger, C.M.,  “Me and we: the interplay between individual and group behavioral variation in social collectives.” Current Opinion in Insect Science. 5: 16-24 (2014). (pdf)

Muli, E., Patch, H., Frazier, M., Frazier, J., Torto, B., Baumgarten, T., Kilonzo, J., Mumoki, F., Masiga, D., Tumlinson, J. and Grozinger, C. “Evaluation of the distribution and impacts of parasites, pathogens, and pesticides on honey bee (Apis mellifera) populations in East Africa.” PloS One. 9(4): p.e94459 (2014). (link)

Manfredini, F., Lucas, C., Nicolas, M., Keller, L., Shoemaker, D. and Grozinger, C.M. “Molecular and social regulation of worker division of labour in fire ants.” Molecular Ecology. 23(3): 660-672 (2014). (link)

Ostiguy, N. and Eitzer, B. “Overwintered brood comb honey: colony exposure to pesticide residues.” Journal of Apicultural Research. 53(3): 413-421 (2014). (link)

Russo, L., Memmott, J., Montoya, D., Shea, K. and Buckley, Y.M. “Patterns of introduced species interactions affect multiple aspects of network structure in plant–pollinator communities.” Ecology. 95(10): 2953-2963 (2014). (link)

LaBar, T., Campbell, C., Yang, S., Albert, R. and Shea, K. “Restoration of plant–pollinator interaction networks via species translocation.” Theoretical ecology. 7(2): 209-220 (2014). (link)

Schmehl, D. R., P. E. A. Teal, et al. (2014). "Genomic analysis of the interaction between pesticide exposure and nutrition in honey bees (Apis mellifera)." Journal of Insect Physiology 71(0): 177-190.

Zhu, W., Schmehl, D.R., Mullin, C.A., Frazier, J.L.  "Four common pesticides, their mixtures and a formulation solvent in the hive environment have high oral toxicity to honey bee larvae" PLoS One. 9(1):e77547 (2014) (PSU news story)

Chen J, Mullin CA.  "Determination of nonylphenol ethoxylate and octylphenol ethoxylate surfactants in beehive samples by high performance liquid chromatography coupled to mass spectrometry" Food Chem. 158:473-9 (2014).

Chejanovsky, N., Ophir, R., Schwager, M.S., Slabezki, Y., Grossman, S., Cox-Foster, D. "Characterization of viral siRNA populations in honey bee colony collapse disorder" Virology. 454-455:176-83 (2014).

Shutler, D., Head, K., Burgher-MacLellan, K.L., Colwell, M.J., Levitt, A.L., Ostiguy, N., Williams, G.R. "Honey Bee Apis mellifera Parasites in the Absence of Nosema ceranae Fungi and Varroa destructor Mites" PLoS One 9(6):e98599 (2014). (PSU news story)

Amsalem, E., Teal, P., Grozinger, C.M., and A. Hefetz.  "Precocene-I inhibits juvenile hormone biosynthesis, ovarian activation, aggression and alters sterility signal production in bumble bee (Bombus terrestris) workers" Journal of Experimental Biology (2014, in press).

Vaudo, A.D., Patch, H.M., Mortensen, D.A., Grozinger, C. M., and J. F. Tooker. "Bumble bees exhibit daily behavioral patterns in pollen foraging".  Arthropod-Plant Interactions 8(4): 273-283 (2014).

Muli*, E., Patch*, H.M., Frazier*, M., Frazier, J., Torto, B., Baumgarten, T., Kilonzo, J., Kilmani, J., Mumoki, F., Masiga, D., Tumlinson, J., and C.M. Grozinger. "Evaluation of distribution and impacts of parasites, pathogens, and pesticides on honey bee (Apis mellifera) populations in East Africa" PLoS ONE 9(4): e94459 (2014). (PSU news story)

Amsalem, E., Malka, O., Grozinger, C.M., and A. Hefetz.  "Exploring the role of juvenile hormone and vitellogenin in reproduction and social behavior in bumble bees" BMC Evolutionary Biology 14:45 (2014).

Malka, O., Niño, E.L., Grozinger, C.M., and A. Hefetz.  “Genomic analysis of the interactions between social environment and social communication systems in honey bees (Apis mellifera)” Insect Biochemistry and Molecular Biology 47C: 36-45 (2014).

Toth, A.L., Tooker, J.F., Radhakrishnan, S., Minard, R., Henshaw, M.T., and C.M. Grozinger.  “Shared genes related to aggression, rather than chemical communication, are associated with reproductive dominance in paper wasps (Polistes metricus)”  BMC Genomics 15(1): 75 (2014). (PSU news story)

Cabrera, A.R., Shirk, P.D., Grozinger. C.M., Evans, J.D., and P. A. Teal. "Examining the role of foraging and malvolio in host-finding behavior in the honey bee parasite, Varroa destructor (Anderson & Trueman)".  Archives of Insect Physiology and Biochemistry 85(2): 61-75 (2014).  

Grozinger, C.M., Richards, J., and H. Mattila.  "From molecules to societies: the mechanisms regulating swarming behavior in honey bees (Apis spp)" Apidologie 45:237-346 (2014).

Shapiro, L.R., Seidl-Adams, I., De Moraes, C.M., Stephenson, A.G., Mescher, M.C. "Dynamics of short- and long-term association between a bacterial plant pathogen and its arthropod vector" Sci Rep. 4:4155 (2014)

Kariyat, R.R., Scanlon, S.R., Moraski, R.P., Stephenson, A.G., Mescher, M.C., De Moraes, C.M. "Plant inbreeding and prior herbivory influence the attraction of caterpillars (Manduca sexta) to odors of the host plant Solanum carolinense (Solanaceae)" Am J Bot. 101(2):376-80 (2014)


Biddinger, D.J., Robertson, J.L., Mullin, C., Frazier, J., Ashcraft, S.A., Rajotte, E.G., Joshi, N.K., Vaughn, M.  "Comparative toxicities and synergism of apple orchard pesticides to Apis mellifera (L.) and Osmia cornifrons (Radoszkowski)"  PLoS One. 8(9):e72587 (2013).

Russo L, Debarros N, Yang S, Shea K, Mortensen D. " Supporting crop pollinators with floral resources: network-based phenological matching." Ecol Evol. 3(9):3125-40 (2013).

Levitt, A.L., Singh, R., Cox-Foster, D.L., Rajotte, E., Hoover, K., Ostiguy, N., Holmes, E.C. "Cross-species transmission of honey bee viruses in associated arthropods." Virus Research 176(1-2):232-40 (2013).

Supple, M.A., Hines, H.M., Dasmahapatra, K.K., Lewis, J.J., Nielsen, D.M., Lavoie, C., Ray, D.A., Salazar, C., McMillan, W.O., Counterman, B.A. "Genomic architecture of adaptive color pattern divergence and convergence in Heliconius butterflies"  Genome Res. 23(8):1248-57 (2013)

Holt, H.L., Aronstein, K. and C.M. Grozinger. “Chronic parasitization by the microsporidian Nosema causes global expression changes in core nutritional, metabolic, and behavioral pathways in honey bee workers (Apis mellifera)” BMC Genomics 14: 799 (2013). 

Niño, E.L., Malka, O., Hefetz, A. Tarpy, D.R., and C.M. Grozinger.  Chemical profiles of two pheromone glands are differentially regulated by distinct mating factors in honey bee queens (Apis mellifera L.)" PLoS ONE 8(11): e78637 (2013). (PSU news story)

Chen, J., Mullin, C.A. "Quantitative determination of trisiloxane surfactants in beehive environments based on liquid chromatography coupled to mass spectrometry" Environ Sci Technol. 47(16):9317-23 (2013)

Carbera, A.R., Shirk, P.D., Duehl, A.J., Donohue, K.V., Grozinger, C.M., Evans, J.D., and P.E.A. Teal.  “Genomic organization and reproductive regulation of a large lipid transfer protein in the varroa mite, Varroa destructor (Anderson & Trueman)” Insect Molecular Biology 22(5): 505-522 (2013).

Weiner, S.A., Galbraith, D.A., Adams, D.C., Valenzuela, N., Noll, F.B., Grozinger, C.M. and A. L. Toth.  "A survey of DNA methylation across social insect species, life stages, and castes reveals abundant and caste-associated methylation in a primitively social wasp"  Naturwissenschaften 100(8): 795-799 (2013).

Niño, E.L., Tarpy, D.R., and C.M. Grozinger. “Differential effects of insemination volume and substance on reproductive changes in honey bee queens (Apis mellifera L.)” Insect Molecular Biology  22(3): 233-244 (2013).

Nunes, F.M.F., Silva, A., Barchuk, A.R., Bomtorin, A.D., Grozinger, C.M. and Z.L.P. Simoes.  “Non-target effects of double-stranded RNA constructs used in honey bee RNAi assays” Insects 4(1): 90-103 (2013).

Peso, M., Niño, E.L., Grozinger, C.M. and A. B. Barron. “Effect of honey bee queen mating condition on worker ovary activation” Insectes Sociaux 60(2): 123-133 (2013).

Shapiro, L.R., Salvaudon, L., Mauck, K.E., Pulido, H., De Moraes, C.M., Stephenson, A.G., Mescher, M.C. "Disease interactions in a shared host plant: effects of pre-existing viral infection on cucurbit plant defense responses and resistance to bacterial wilt disease" PLoS One. 8(10):e77393 (2013).  

Simmons, H.E., Dunham, J.P., Zinn, K.E., Munkvold, G.P., Holmes, E.C., Stephenson, A.G. "Zucchini yellow mosaic virus (ZYMV, Potyvirus): vertical transmission, seed infection and cryptic infections"  Virus Res. 176(1-2):259-64 (2013).

Kariyat, R.R., Balogh, C.M., Moraski, R.P., De Moraes, C.M., Mescher, M.C., Stephenson, A.G. "Constitutive and herbivore-induced structural defenses are compromised by inbreeding in Solanum carolinense (Solanaceae)" Am J Bot. 100(6):1014-21 (2013).

Kariyat, R.R., Mauck, K.E., Balogh, C.M., Stephenson, A.G., Mescher, M.C., De Moraes, C.M. "Inbreeding in horsenettle (Solanum carolinense) alters night-time volatile emissions that guide oviposition by Manduca sexta moths" Proc Biol Sci. 280(1757):20130020 (2013).


Duennes, M.A., Lozier, J.D., Hines, H.M., and S. A. Cameron. “Geographical patterns of genetic divergence in the widespread Mesoamerican bumble bee Bombus ephippiatus (Hymenoptera: Apidae)”. Molecular Phylogenetics and Evolution 64(1):219-31 (2012). doi: 10.1016/j.ympev.2012.03.018. (link)

Hines, H.M., Papa, R., Ruiz, M., Papanicolaou, A., Wang, C., Nijhout. H.F., McMillan, W.O., and R. D. Reed. “Transcriptome analysis reveals novel patterning and pigmentation genes underlying Heliconius butterfly wing pattern variation”. BMC Genomics 13:288 (2012). doi: 10.1186/1471-2164-13-288. (link)

Campbell, C., Yang, S., Shea, K., and R. Albert. “Topology of plant-pollinator networks that are vulnerable to collapse from species extinction.” Physical Review. E, Statistical, Nonlinear and Soft Matter Physics 86(2):021924 (2012) doi: 10.1103/PhysRevE.86.021924 (link)

Cardoza, Y.J., Harris, G.K. and C.M. Grozinger.  “Effects of soil quality enhancement on pollinator-plant interactions” Psyche 2012: 581458, 8p (2012). doi:10.1155/2012/581458 (link)

Niño, E.L., Malka, O., Hefetz, A., Teal, P., Hayes, J. and C.M. Grozinger. “Effects of honey bee queen (Hymenoptera: Apis mellifera) insemination volume on queen-worker interactions” Journal of Insect Physiology 58(8):1082-1089 (2012). doi: 10.1016/j.jinsphys.2012.04.015 (link)

Wang, Y., Kocher, S.D., Linksvayer T.A., Grozinger, C.M., Page, R.E., and G.V. Amdam. “Regulation of behaviorally-associated gene pathways in worker honey bee ovaries” Journal of Experimental Biology 215(1): 124-134 (2012). doi:10.1242/jeb.060889 (link)

Kariyat, R.R., De Moraes, C.M., Stephenson, A.G., and M.C. Mescher. “Inbreeding increases susceptibility to powdery mildew (Oidium neolycopersici) infestation in horsenettle (Solanum carolinense L)” Plant Signaling and Behavior 7:1-4 (2012). doi: 10.4161/psb.20602 (link)

Shapiro, L., C.M. De Moraes, A.G. Stephenson and M.C. Mescher. “Pathogen effects on vegetative and floral odors mediate vector attraction and host exposure in a complex pathosystem” Ecology Letters. 2012. doi: 10.1111/ele.12001 (link)

Simmons, H.E., Dunham, J.P., Stack, J.C., Dickins, B.J.A., Pagan, I., Holmes, E.C., and A.G. Stephenson. “Deep sequencing reveals persistence of intra- and inter-host genetic diversity in natural and greenhouse populations of Zucchini yellow mosaic virus” Journal of General Virology 93:1831-1840 (2012).  doi: 10.1099/vir.0.042622-0 (link)

Kariyat, R.R., Mena-Ali, J.I., Forry, B., Mescher, M.C., De Moraes, C.M., and A.G. Stephenson. “Inbreeding, herbivory, and the transcriptome of Solanum carolinense L” Entomologia Experimentalis et Applicata. 144: 134–144 (2012). doi: 10.1111/j.1570-7458.2012.01269.x (link)

Kariyat, R.R., Mauck, K.E., De Moraes, C.M., Stephenson, A.G. and M.C. Mescher. “Inbreeding alters volatile signaling phenotypes and influences tri-trophic interactions in horsenettle (Solanum carolinense L)” Ecology Letters 15: 301–309. doi: 10.1111/j.1461-0248.2011.01738.x (link)

Stephenson, A.G. “Safe sex in plants” New Phytologist 193:827-829 (2012). doi: 10.1111/j.1469-8137.2011.04035.x (link)

Ciarlo, T.J., Mullin, C.A., Frazier, J.L., and D.R. Schmehl. “Learning impairment in honey bees caused by agricultural spray adjuvants” PLoS ONE 7: 1-12 e40848 (2012).  doi: 10.1371/journal.pone.0040848 (link)

Chen, J., Frazier, J.L., Frazier, M.T., and C. A. Mullin. “Identification of organosiloxane surfactants in agrochemical spray adjuvants using liquid chromatography coupled to mass spectrometry”  In: American Bee Research Conference, Riverdale, MD. American Bee Journal 152(4): 400 (2012). (Abstract)

Freiberg M., De Jong, D., Message, D., and D. Cox-Foster D. “First report of sacbrood virus in honey bee (Apis mellifera) colonies in Brazil”. Genetics and  Molecular Reserch 11(3):3310-4 (2012). Doi: 10.4238/2012.September.12.14  (link)

Johnson, R. M., Ellis, M. D., Mullin, C. A., and M. Frazier. “Pesticides and honey bee toxicity in the United States”.  Honey Bee Colony Health - Challenges and Sustainable Solutions (D. Sammataro, and J. A. Yoder, Eds.), pp. 145-160 (2012). CRC Press, Boca Raton, FL.

Drummond, F, Aronstein, K., Chen, J., Ellis, J., Evans, J., Ostiguy, N., Sheppard, W., Spivak, M., and K. Visscher.  “First two years of the stationary hive project.”   Bee Culture April  2012. (link)

Richard, F.J., Holt, H.L., and C.M. Grozinger. “Effects of immunostimulation on social behavior, chemical communication and genome-wide gene expression in honey bee workers (Apis mellifera).” BMC Genomics. 13:558 (2012).  doi: 10.1186/1471-2164-13-558  (link)

Zhu, W., T. Reluga, C. Mullin and J. Frazier.  “A stage-structured model of honey bee colony population dynamics assessing impacts of pesticides and other stressors.”  In: American Bee Research Conference, Riverdale, MD. American Bee Journal 152(4): 407-408 (2012). (Abstract)

Niño, E. L., D. R. Tarpy, and C. M. Grozinger. “Differential effects of insemination volume and substance on reproductive changes in honey bee queens (Apis mellifera L.)” Insect Molecular Biology (In press)

Harth, J.E., Deveney, T.W., Winsor, J.A., Ferrari, M.J., and A.G. Stephenson. “Background and development of a model to predict the fate of an escaped virus resistant transgene from Cucurbita pepo”. Proceedings of the Xth Eucarpia Conference: Cucurbitaceae (In press).


Kocher S.D. and C. M. Grozinger. “Cooperation, conflict and the evolution of queen pheromones” Journal of Chemical Ecology 37(11):1263-1275 (2011). doi: 10.1007/s10886-011-0036-z (link)

Richard, F.J., Schal, C. Tarpy, D.R., and C.M. Grozinger. “Effects of insemination number on honey bee queen Dufour’s gland secretion” Journal of Chemical Ecology 37(9):1027-1036 (2011). doi: 10.1007/s10886-011-9999-z (link)

Fussnecker, B.L., McKenzie, A.M., and C.M. Grozinger. “The role of cGMP in modulating behavior, physiology, and brain gene expression in response to queen mandibular pheromone in honey bees” Journal of Comparative Physiology A 197(9): 939-948 (2011). doi: 10.1007/s00359-011-0654-5 (link)

Bloch, G. and C.M. Grozinger. “Social molecular pathways and the evolution of bee societies” Philosophical Transactions of the Royal Society B 366(1574): 2155-2170 (2011). doi: 10.1098/rstb.2010.0346 (link)

Niño, E.L., Tarpy, D.R., and C.M. Grozinger. “Genome-wide analysis of brain transcriptional changes in honey bee (Apis mellifera L.) queens exposed to carbon dioxide and physical manipulation” Insect Molecular Biology 20(3): 387-398 (2011). doi: 10.1111/j.1365-2583.2011.01072.x (link)

Kariyat, R.R., Scanlon, S.C., Mescher, M.C., De Moraes, C.M., and A.G. Stephenson. “Inbreeding depression in Solanum carolinense (Solanaceae) under field conditions and implications for mating system evolution” PLoS One 6(12): e28459 (2011) doi:10.1371/journal.pone.0028459 (link)

Simmons, H.E., Holmes, E.C., Gildow, F.E., Bothe-Goralczyk, M.A., and A.G. Stephenson. “Experimental verification of seed transmission in Zucchini yellow mosaic virus” Plant Disease 95:751-754 (2011). doi: 10.1094/PDIS-11-10-0843 (link)

Simmons, H.E., Holmes, E.C., and A.G. Stephenson. “Rapid turnover of intra-host genetic diversity in Zucchini yellow mosaic virus” Virus Research 155:389-396 (2011). doi: 10.1016/j.virusres.2010.11.007 (link)


Cornman, S., Schatz, M.C., Johnston, J.S., Chen, Y-P., Pettis, J., Hunt, G., Bourgeois, L., Elsik, C. Anderson, D. Grozinger, C.M. and J. D. Evans. “Genomic survey of the ectoparasitic mite Varroa destructor, a major pest of the honey bee Apis mellifera” BMC Genomics 11:602; 15p (2010). doi: (link)

Kocher, S.D., Richard, F.J., Tarpy, D.R., and C.M. Grozinger.  "The effects of mating and instrumental insemination on queen honey bee flight behaviour and gene expression" Insect Molecular Biology 19(2): 153-162 (2010). doi: 10.1111/j.1365-2583.2009.00965.x (link)

Wilson, E. E., Sidhu, C.S., LeVan, K.E., and D. A. Holway. "Pollen foraging behavior of solitary Hawaiian bees revealed through molecular pollen analysis" Molecular Ecology DOI: 10.1111/j.1365-294X.2010.04849.x (2010) (link)

Shpigler, H., Patch, H.M., Cohen, M., Fan, Y., Grozinger, C.M., and G. Bloch. “A gene for queen control: Krüppel homolog 1 is linked to social organization in bees” BMC Evolutionary Biology 10:120 (2010). (link)

Biddinger, D. J., H. Ngugi, J. Frazier, M. Frazier, T. Leslie, and L. R. Donovall.  2010.  Development of the mason bee, Osmia cornifrons, as and alternative pollinator to honey bees and as a targeted delivery system for biological control agents in the management of fire blight.  Penn Fruit News, vol. 90 (2): 35-44. (pdf)

Donovall, L. R and D. vanEngelsdorp. "A Checklist of the Bees (Hymenoptera: Apoidea) of Pennsylvania." Journal of the Kansas Entomological Society 83(1): 7-24 (2010). (pdf)

Mullin,C., Frazier, M., Frazier J., Ashcraft S., Simonds, R., vanEngelsdorp, D., Pettis, J. "High Levels of Miticides and Agrochemicals in North American Apiaries: Implications for Honey Bee Health" PLoS ONE 5(3): e9754 (2010). (pdf)

Kocher, S.D., Ayroles, J.F., Stone, E.A.  and C.M. Grozinger. “Individual variation in pheromone response correlates with reproductive traits and brain gene expression in worker honey bees” PLoS ONE 5(2): e9116 (2010). (link)

Fan, Y., Richard, FJ., Rouf, C. and C.M. Grozinger. “Effects of queen mandibular pheromone on nestmate recognition in worker honey bees (Apis mellifera)” Animal Behavior 79(3): 649-656 (2010). (link)

Williams, Geoffrey R., Richard E.L. Rogers, Abby L. Kalkstein, Benjamin A. Taylor, Dave Shutler, Nancy Ostiguy. Deformed wing virus in western honey bees (Apis mellifera) from Atlantic Canada and the first description of an overtly-infected emerging queen. Journal of Invertebrate Pathology. 101(1) 77-79. (2010)

Ostiguy, N.  Managed Pollinator CAP Coordinated Agricultural Project A National Research and Extension Initiative to Reverse Pollinator Decline Sustainable Beekeeping. American Bee Journal. 150 (2):149-152 (2010)

vanEngelsdorp, D., and M. D. Meixner. “A Historical Review of Managed Honey Bee Populations in Europe and the United States and the Factors That May Affect Them” Journal of Invertebrate Pathology 103:S80–S95 (2010) (link)

vanEngelsdorp, D., J. Hayes Jr, R. M. Underwood, and J. S. Pettis. 2010. A survey of honey bee colony losses in the United States, fall 2008 to spring 2009. Journal of Apicultural Research 49 7 -14. (pdf)

Singh R., Levitt, A.L., Rajotte, E.G., Holmes, E.C., Ostiguy, N., vanEngelsdorp D., Lipkin, W.I., Depamphilis, C.W., Toth, A.L., and D. Cox-Foster. “RNA viruses in hymenopteran pollinators: evidence of inter-Taxa virus transmission via pollen and potential impact on non-Apis hymenopteran species” PLoS ONE 5(12):e14357 (2010). doi: (link)

Hunter. W., Ellis. J., vanEngelsdorp, D., Hayes, J., Westervelt, D., Glick, E., Williams, M., Sela, I., Maori, E., Pettis, J., Cox-Foster, D., and N. Paldi. “Large-scale field application of RNAi technology reducing Israeli acute paralysis virus disease in honey bees (Apis mellifera, Hymenoptera: Apidae)” PLoS Pathogens 6(12): e1001160 (2010). doi: 10.1371/journal.ppat.1001160 (link)


Williams, G.R., Rogers, R.E.L., Kalkstein, A.L., Taylor, B.A., Shutler D, and N. Ostiguy. “Deformed wing virus in western honey bees (Apis mellifera) from Atlantic Canada and the first description of an overtly-infected emerging queen” Journal of Invertebrate Pathology 101(1) 77-79 (2009). doi: 10.1016/j.jip.2009.01.004 (link)

Biddinger, D. J., H. Ngugi, & J. Frazier.  2009.  Development of the Mason Bee, Osmia cornifrons, as a Targeted Delivery System for Biocontrol Agents in the Management of Fire Blight.  Penn Fruit News, vol. 89 (2): 95-100. (pdf)

Biddinger, D., J. Frazier, M. Frazier, E. Rajotte, L. Donovall, and T. Leslie.  2009.  Solitary bees as alternative pollinators in Pennsylvania fruit crops.  Penn Fruit News, vol. 89 (2): 84-94. (pdf)

Patch, HM; Velarde, RA; Walden, KKO; Robertson, HM.  A Candidate Pheromone Receptor and Two Odorant Receptors of the Hawkmoth Manduca sexta. Chemical Senses  34 (4):305-316 (2009)

Kocher, S.D., Richard, F.J., Tarpy, D.R., and C.M. Grozinger.  “Queen reproductive state modulates queen pheromone production and queen-worker interactions in honey bees” Behavioral Ecology 20: 1007-1014 (2009)

Kocher, S.D., Richard, F.J., Tarpy, D.R., and C.M. Grozinger.  "The effects of mating and instrumental insemination on honey bee behavior, physiology, and brain gene expression" Insect Molecular Biology Published online December 11, 2009, doi: 10.1111/j.1365-2583.2009.00965.x.

vanengelsdorp, D., and L. R. Donaval. 2009. Note: New Plant Host Record and First Record of the Burrower Bug Sehirus cinetus (Palisot de Beauvois) (Hemiptera: Cydnidae) Associating with Honey Bees, Apis mellifera L. (Hymenoptera: Apidae). Proceedings of the Entomology Society of Washington 111: 903-906. (pdf)

vanEngelsdorp, D., J. D. Evans, L. Donovall, C. Mullin, M. Frazier, J. Frazier, D. R. Tarpy, J. Hayes Jr, and J. S. Pettis. 2009. "Entombed Pollen": A new condition in honey bee colonies associated with increased risk of colony mortality. Journal of Invertebrate Pathology 101: 147-149. (pdf)

vanEngelsdorp, D., J. D. Evans, C. Saegerman, C. Mullin, E. Haubruge, B. K. Nguyen, M. Frazier, J. Frazier, D. Cox-Foster, Y. Chen, R. Underwood, D. R. Tarpy, and J. S. Pettis. 2009. Colony Collapse Disorder: A Descriptive Study. PLoS ONE, 4: e6481. (Link)

vanEngelsdorp, D., S. Gebauer, R. Underwood. 2009. A modified Two-queen System: “Tower” Colonies Allowing for Easy Drone Brood Removal for Varroa Mite Control.  Science of Bee Culture 1(1): 1-4. (pdf)

Cox-Foster, D., and D. vanEngelsdorp. 2009. Solving the mystery of the disappearing bees. Scientific American: April 40-47. (Link to article)

vanengelsdorp, D., and L. R. Donaval. 2009. Note: New Plant Host Record and First Record of the Burrower Bug Sehirus cinetus (Palisot de Beauvois) (Hemiptera: Cydnidae) Associating with Honey Bees, Apis mellifera L. (Hymenoptera: Apidae). Proceedings of the Entomology Society of Washington 111: 903-906. (pdf)

Kocher, S.D., Richard, F.J., Tarpy, D.R., and C.M. Grozinger.  “Queen reproductive state modulates queen pheromone production and queen-worker interactions in honey bees” Behavioral Ecology 20(5):1007-1014;

Alaux, C., Y. Le Conte, H. Heather, S. Rodrigues-Zas, C.M. Grozinger, S. Sinha, and G. E. Robinson. “Regulation of brain gene expression in honey bees by brood pheromone” Genes, Brain, and Behavior. 8(3):309-19 (2009)


vanEngelsdorp D. R. Underwood, and D. Cox Foster 2008 Flash fumigation of honey bee Apis mellifera (Hymenoptera: Apidae) colonies with formic and acetic acids for the control of Varroa destructor (Acari: Varroidae). Journal of Economic Entomology101(2) 256-264 (pdf)

G Palacios, J Hui, P L Quan, A Kalkstein, K S Honkavuori, A V Bussetti, S Conlan, J Evans, Y P Chen, D vanEngelsdorp, H Efrat, J Pettis, D Cox-Foster, E C Holmes, T Briese, W I Lipkin 2008 Genetic analysis of Israel Acute Paralysis Virus: distinct clusters are circulating in the United States. Journal of Virology 23: JVI.00251-08 (pdf)

vanEngelsdorp, D., J. Hayes, Jr., R. M. Underwood, and J. Pettis. 2008. A Survey of Honey Bee Colony Losses in the U.S., Fall 2007 to Spring 2008. PLoS ONE 3: e4071. (Link)

Richard, F.J., A. Aubert, and C.M. Grozinger. “Modulation of nestmate recognition by immune stimulation in honey bees, Apis mellifera”.  BMC Biology 6:50 (2008).

Fussnecker, B. and C.M. Grozinger. “Dissecting the role of Kr-h1 brain gene expression in foraging behavior in honey bees (Apis mellifera)”.  Insect Molecular Biology. 17(5):515-22 (2008).

Kocher, S.D.,Richard, F.J., Tarpy, D.R., and C.M. Grozinger. “Genomic analysis of post-mating changes in the honey bee queen”. BMC Genomics. 9(1):232 (2008).

Fischer, P. and Grozinger. “Pheromonal regulation of starvation resistance in honey bee workers”. Naturwissenschaften. 95(8):723-9 (2008).


Yang, X; Cox-Foster, D.  Effects of parasitization by Varroa destructor on survivorship and physiological traits of Apis mellifera in correlation with viral incidence and microbial challenge.  Parasitology 134405-412 (2007)

Ostiguy, N., D. Cox-Foster, A. Kalkstein and O. Thompson. The Continuing Story of Honey Bee Viruses. American Bee Journal 147(5):441 (2007)

Underwood, R., D. vanEngelsdorp 2007 Colony Collapse Disorder: Have we seen this Before? Bee Culture 135(7) 13-18 (pdf)

vanEngelsdorp, D., R. Underwood, D. Caron, J. Hayes, Jr. 2007 An Estimate of Managed Colony Losses in the Winter of 2006 - 2007: A Report Commissioned by the Apiary Inspectors of America. American Bee Journal 147 (7) 599-603. (pdf)

Arbogast, Richard T., Baldwyn Torto, Dennis vanEngelsdorp, and Peter E. A. Teal 2007 An Effective Trap and Bait combination for monitoring the Small Hive Beetle Aethina Tumida (Coleoptera: Nitidulidae) Florida Entomologist 90 (2) 404-406. (pdf)

Torto, Baldwyn, Richard T. Arbogast, Dennis vanEngelsdorp, Steven Willms, Dusti Purcell, Drion Boucias, James H. Tumlinson, Peter E. A. Teal. 2007 Trapping of Aethina tumida Murry (Coleoptera: Nitidulidae) from Apis mellifera L. (hymenoptera: Apidae) Colonies with an In-Hive Baited Trap. Environmental Entomology 36(5):1018-1024. (pdf)

Cox-Foster D.L., Conlan S., Holmes E.C, Palacios G., Evans J.D., Moran N.A, Quan P-L., Briese T., Geiser D.M., Martinson V., vanEngelsdorp D., Kalkstein, A.L., Drysdale A, Hui J., Zhai J., Cui L, Hutchinson S, Simons J.F., Egholm M, Pettis J.S., Lipkin W.I.  2007. A metagenomic survey of microbes in honey bee colony collapse disorder. Science  318: 283-286. (pdf)

Torto, B, R. T. Arbogast, H. Alborn, A. Suazo, D, vanEngelsodorp, D. Boucias, J. T. Tumlinson, P. E. A. Teal. 2007. Composition of volatiles from fermenting pollen dough and attractiveness to small hive beetle Aethina tumida a parasite of the honeybee Apis mellifera. Apidologie 38, 380-389. (Link)

Richard, F.J., Tarpy, D.R, and C.M. Grozinger. “Effects of insemination quantity on honey bee queen physiology”. PLoS ONE , 2(10):e980 (2007).

Grozinger, C.M., Fan, Y., Hoover, S.E.R.  and M.L. Winston. “Genome-wide analysis reveals differences in brain gene expression patterns associated with caste and reproductive status in honey bees (Apis mellifera).” Molecular Ecology 16(22):4837-48 (2007).

Grozinger, C.M., Fischer, P, and J.E. Hampton.  “Uncoupling primer and releaser responses to pheromone in honey bees.” Naturwissenschaften 94(5):375-9. (2007)

Grozinger, C.M. and Robinson, G.E. “Endocrine modulation of a pheromone responsive gene in the honey bee brain” Journal of Comparative Biology A 193(4):461-70 (2007).


Kevan, PG; Hannan, MA; Ostiguy, N; Guzman-Novoa, E. A summary of the Varroa-virus disease complex in honey bees.  American Bee Journal.  146 (8):694-697 (2006)

Tooker, J. F., M. Hauser, and L. M. Hanks. 2006. Floral Host Plants of
Syrphidae and Tachinidae (Diptera) of Central Illinois. Annals of the
Entomological Society of America 99: 96-112.


Shen, MQ; Cui, LW; Ostiguy, N; Cox-Foster, D.  Intricate transmission routes and interactions between picorna-like viruses (Kashmir bee virus and sacbrood virus) with the honeybee host and the parasitic varroa mite.  Journal of General Virology 862281-2289 Part 8 (2005)

Shen, MQ; Yang, XL; Cox-Foster, D; Cui, LW.  The role of varroa mites in infections of Kashmir bee virus (KBV) and deformed wing virus (DWV) in honey bees. Virology  342 (1):141-149 (2005)


Sammataro, D; Hoffman, GD; Wardell, G; Finley, J; Ostiguy, N.  Testing of a combination of control tactics to manage Varroa destructor (Acari : Varroidae) population levels in honey bee (Hymenoptera : Apidae).  International Journal of Acarology  30 (1):71-76 (2004)


Tooker, J. F., P. F. Reagel, and L. M. Hanks. 2002. Nectar sources of
day-flying Lepidoptera of central Illinois. Annals of the
Entomological Society of America 95: 84-96.


Tooker, J. F., and L. M. Hanks. 2000. Flowering plant hosts of adult
hymenopteran parasitoids of central Illinois. Annals of the
Entomological Society of America 93: 580-588.