lab members in bold


McNeil, D.J., E. McCormick, A.C. Heimann, M. Kammerer, M.R. Douglas, S.C. Goslee, C.M. Grozinger, H.M. Hines. 2020. Bumble bees in landscapes with abundant floral resources have lower pathogen loads. Nature Scientific Reports 10,22306 (2020).

Rahman, S.R., J. Cnaani, L.N. Kinch, N.V. Grishin, H.M. Hines. A combined RAD-Seq and WGS approach reveals the genomic basis of yellow color variation in bumble bee Bombus terrestrisPREPRINT, bioRxiv. doi:

Villalona, E., Ezray, B.D., Laveaga, E., Agrawal, A.A., Ali, J.G., and H.M. Hines. 2020. The role of toxic nectar secondary compounds in driving differential bumble bee preferences for milkweed flowers. Oecologia

Sharanowski, B.J., R. Ridenbaugh, P.K. Piekarski, G.R. Broad, G.R. Burke, A.R. Deans, A.R. Lemmon, E.R. Lemmon, G. Stewart, J.B. Whitfield, and H.M. Hines. Phylogenomics of Icheumonoidea (Hymenoptera) and implications for evolution of parasitism and viral endogenization. Molecular Phylogenetics and Evolution, 107023; bioRxiv, doi:

 Burke, G.R., H.M. Hines, B.J. Sharanowski. 2020. Endogenization from diverse viral ancestors is common and widespread in parasitoid wasps. bioRxiv, doi:

Ghisbain, G., Lozier, J.D., Rahman, S.R., Ezray, B.D., Tian, L., Ulmer, J.M., Heraghty, S., Strange, J.P., Rasmont, P., and H.M. Hines. 2020. Substantial genetic divergence and lack of recent gene flow support cryptic speciation in a color polymorphic bumble bee (Bombus bifarius) species complex. Systematic Entomology,

Vaudo, A. D., Tooker, J.F., Patch, H.M.,  Biddinger, D.J., Coccia, M., Crone, M.K., Fiely, M., Francis, J.S., Hines, H.M., Hodges, M., Jackson, S.W., Michez, D., Mu, J., Russo, L., Safari, M., Treanore, E.D., Vanderplanck, M., Yip, E.,  Leonard, A.S., and C.M. Grozinger. 2020. Pollen protein:lipid macronutrient ratios guide broad patterns of bee species floral preferences. Insects, 2020 (11):132.



Mikó, I., Rahman, S.R., Jones, A.C., Townley, M.A., Gominho, B., Paudel, S., Stupski, D., Hines, H.M., and R.J. Schilder. 2019. From Spinning Silk to Spreading Saliva: Mouthpart Remodeling in Manduca sexta (Lepidoptera: Sphingidae). Insect Systematics and Diversity, Special Collection: Current Techniques in Morphology, 3(6), November 2019, 2,

Ezray, B.D., D.C. Wham, C. Hill, and H.M. Hines. Unsupervised machine learning reveals mimicry complexes in bumble bees occur along a perceptual continuum. Proceedings of the Royal Society of London, B series, 286: 20191501,

Hines, H.M., and S.R. Rahman. 2019. Evolutionary genetics in insect phenotypic radiations: the value of a comparative genomic approach. Current Opinion in Insect Science.

Wham, D., Ezray, B.D., and H.M. Hines. 2019. Measuring perceptual distance of organismal color pattern using the features of deep neural networks. PREPRINT, bioRxiv, August 15, 2019.

Baker, A., J. Heraty, J. Mottern, J. Zhang, H. M. Hines, A. Lemmon, and E. Lemmon. 2019. Countercurrent dispersals of parasitoids and their ant hosts: the phylogenetics and biogeography of the Oraseminae (Hymenoptera: Eucharitidae). Systematic Entomology,

Tian, L., Rahman, S.R., Ezray,  B.D., Franzini, L., Strange, J.P., Lhomme, P., and H.M. Hines. 2019. A homeotic shift late in development drive mimetic color variation in a bumble bee. Proceedings of the National Academy of Sciences, USA.

Mikó, I., Rahman, S.R., Anzaldo, S.S., van de Kamp, T., Parslow, B.A., Tatarnic, N.J., Wetherington, M.T., Anderson, J., Schilder, R.J., Ulmer, J.M., Deans, A.R., and H.M. Hines. 2019. Fat in the Leg: Function of the expanded hind heg in gasteruptiid wasps (Hymenoptera: Gasteruptiidae). Insect Systematics and Diversity, 3(1), 2.

Ezray, B.D., Wham, D., Hill, C., and H.M. Hines. 2019. Mullerian mimicry in bumble bees is a transient continuum. PREPRINT, bioRxiv, 513275.



Dash, S., Rahman, S., Hines, H.M., and W. Feng. 2018. Incremental BLAST: Incremental addition of new sequence databases through e-value correction. PREPRINT, bioRxiv, 476218.

Tian, L. and H.M. Hines 2018. Morphological characterization and staging of bumble bee pupae. PeerJ, 6, e6089.

Lhomme, P. and H.M. Hines 2018. Ecology and Evolution of Cuckoo Bumble Bees. Annals of the Entomological Society of America,

Lhomme, P. and H.M. Hines 2018. Reproductive dominance strategies of insect social parasites. Journal of Chemical Ecology, 44: 838-850.

Brasero, N., Martinet, B., Lecocq, T., Lhomme, P., Biella, P., Valterová, I., Urbanaova, K., Cornalba, M., Hines, H., and P. Rasmont. 2018. The cephalic labial gland secretions of two socially parasitic bumblebees Bombus hyperboreus (Alpinobombus) and Bombus inexspectatus (Thoracobombus) question their inquiline strategy. Insect Science. 25(1), 75-86.



Hines, H.M. 2017. Book Review: Darwin’s Man in Brazil: The Evolving Science of Fritz Mueller, by David A. West. American Entomologist. 63(1):60.

Harpur, B.A., Dey, A., Albert, J. R., Patel, S., Hines, H.M., Hasselmann, M., Packer, L. Zayed, A. 2017. Queens and Workers Contribute Differently to Adaptive Evolution in Bumble Bees and Honey Bees. Genome Biology and Evolution, 9(9), 2395–2402.

Hines, H.M., Witkowski, P., Wilson, J.S., Wakamatsu, K. 2017. Melanic variation underlies aposematic color variation in two hymenopteran mimicry systems. PLOS ONE 12(7): e0182135.

Van Belleghem, S.M., Rastas, P., Papanicolaou, A., Martin, S.H., Arias, C.F., Supple, M.A., Hanly, J.J., Mallet, J., Lewis, J.J., Hines, H.M., Ruiz, M., Salazar, C., Linares, M., Moreira, G.R.P., Jiggins, C.D., Counterman, B.A., McMillan, W.O., and R. Papa. 2017. Complex modular architecture around a simple toolkit of wing pattern genes. Nature Ecology & Evolution 1: p.0052.



Mikó I, Trietsch C, Sandall EL, Yoder MJ, Hines H, Deans AR. 2016. Malagasy Conostigmus (Hymenoptera: Ceraphronoidea) and the secret of scutes. PeerJ 4:e2682

Nadeau, N.J., C. Pardo-Diaz, A. Whibley, M.A. Supple, S.V. Saenko, R.W.R. Wallbank, G.C. Wu, L. Maroja, L. Ferguson, J.J. Hanly, H. Hines, C. Salazar, R. M. Merrill, A. J. Dowling, R. H. ffrench-Constant, V. Llaurens, M. Joron, W. O. McMillan, C. Jiggins. 2016. The gene cortex controls mimicry and crypsis in butterflies and moths. Nature 543: 106-110.

Nixon, A.E., H. Hines. 2016. Bumblebee thermoregulation: Understanding the thermal properties of physical features of bumble bees. The Penn State McNair Journal. 21: 99-115.



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

Deans, A.R., Lewis, S.E., Huala, E., <70 alphabetical authors including H.M.Hines>, and P. Mabee. 2015. Finding our way through phenotypes. PLoS Biology 13(1): e1002033 doi: 10.1371/journal.pbio.1002033

Condamine, F. and H. Hines, 2015. Historical species losses in bumblebee evolution. Biology Letters, 11: 20141049.



Supple, M. A., H. M. Hines, K. K. Dasmahapatra, J. J. Lewis, D. M. Nielsen, C. Lavoie, D. A. Ray, C. Salazar, W. O. McMillan, and B. A. Counterman. 2013. Genomic architecture of adaptive color pattern divergence and convergence in Heliconius butterflies. Genome Research, PREPRINT, doi: 10.1101/gr.150615.112

************ Pre-PSU Publications************


Hines, H. M., P. Williams. 2012. Evolution of Müllerian mimicry in the highly color polymorphic bumble bee Bombus trifasciatus and its comimics. Zoological Journal of the Linnean Society 166:805-826.

Hines, H. M.*, R. Papa*, M. Ruiz, A. Papanicoloau, C. Wang, H. F. Nijhout, W. O. McMillan**, and R. Reed**. Transcriptome analysis reveals novel patterning and pigmentation genes underlying Heliconius butterfly wing pattern variation. BMC Genomics 13: 288. *, ** equal contributions

The Heliconius Genome Consortium. 2012. Butterfly genome reveals promiscuous exchange of mimicry adaptations among species. Nature,

Duennes, M. A., Lozier, J. D., Hines, H. M., and S. A. Cameron. 2012. Geographical patterns of genetic divergence in the widespread Mesoamerican bumble bee Bombus ephippiatus (Hymenoptera: Apidae). Molecular Phylogenetics and Evolution 64 (1): 219-231.

Miko, I., Friedrich, F., Yoder, M. J., Hines, H. M., Deitz, L. L., Bertone, M. A., Seltman, K. C., Wallace, M. S., and A. R. Deans. 2012. On dorsal prothoracic appendages in treehoppers (Hemiptera: Membracidae) and the nature of morphological evidence. PLoS ONE 7(1): e30137. doi:10.1371/journal.pone.0030137


Hines, H. M., Counterman, B. A., Papa, R., Albuquerque de Moura, P., Cardoso, M. Z., Linares, M., Mallet, J., Reed, R. D., Jiggins, C. D., Kronforst, M. R., and W. Owen McMillan. 2011. Wing patterning gene redefines the mimetic history of Heliconius butterflies. Proceedings of the National Academy of Sciences, USA 108: 19666-19671.

Reed, R. D., Papa, R., Martin, A., Hines, H. M., Counterman B. A., Pardo-Diaz, C., Jiggins, C. D., Chamberlain, N. L., Kronforst M. R., Chen, R., Halder, G., Nijhout, H. F., and W. O. McMillan. 2011. optix drives the repeated convergent evolution of butterfly wing pattern mimicry. Science 333: 1137-1141.


Hines, H. M. and S. A. Cameron. 2010. The phylogenetic position of the bumble bee inquiline Bombus inexspectatus and implications for the evolution of social parasitism. Insectes Sociaux 57: 379-383.

Counterman, B. A., Arajuo-Perez, F., Hines, H. M., Baxter, S. W., Morrison, C. M., Lindstrom, D. P., Papa, R., Ferguson, L., Joron, M., ffrench-Constant, R., Smith, C., Nielsen, D. M., Chen, R., Jiggins, C. D., Reed, R. D., Halder, G., Mallet, J., and W. O. McMillan. 2010. Genomic hotspots for adaptation: population genetics of Müllerian mimicry in Heliconius erato. PLoS Genetics 6 (2): e10000794.


Williams, P. H., Cameron, S. A., Hines, H. M., Cederberg, B., and P. Rasmont. 2008. A simplified subgeneric classification of the bumblebees (genus Bombus). Apidologie 39: 46-74.

Hines, H. M. 2008. Historical biogeography, divergence times, and diversification patterns of bumble bees (Hymenoptera: Apidae: Bombus). Systematic Biology 57: 58-75.


Cameron, S. A., Hines, H. M., and P. H. Williams. 2007. A comprehensive phylogeny of the bumble bees (Bombus). Biological Journal of the Linnean Society 91: 161-188.

Hines, H. M., Cameron, S. A., and A. R. Deans. 2007a. Nest architecture and foraging behavior in Bombus pullatus (Hymenoptera: Apidae), with comparisons to other tropical bumble bees. Journal of the Kansas Entomological Society 80: 1-15.

Hines, H. M., Hunt, J. H., O’Connor, T. K., Gillespie, J. J., and S. A. Cameron. 2007b. Multigene phylogeny reveals eusociality evolved twice in vespid wasps. Proceedings of the National Academy of Sciences, USA 104: 3295-3299.


Hines, H. M., Cameron, S. A., and P. H. Williams. 2006. Molecular phylogeny of the bumble bee subgenus Pyrobombus (Hymenoptera: Apidae: Bombus) with insights into gene utility for lower-level analysis. Invertebrate Systematics 20: 289-303.


Rasmont, P., Terzo, M., Aytekin, A. M., Hines, H., Urbanova, K., Cahlikova, L., and I. Valterova. 2005. Cephalic secretions of the bumblebee subgenus Sibiricobombus Vogt suggest Bombus niveatus Kriechbaumer and Bombus vorticosus Gerstaecker are conspecific (Hymenoptera, Apidae, Bombus). Apidologie 36: 571-584.

Hines, H. M., and S. D. Hendrix. 2005. Bumble bee (Hymenoptera: Apidae) diversity and abundance in tallgrass prairie patches: Effects of local and landscape floral resources. Environmental Entomology 34: 1477-1484.


Nelson, D. R., Hines, H., and B. Stay. 2004. Methyl-branched hydrocarbons, major components of the waxy material coating the embryos of the viviparous cockroach Diploptera punctata. Comparative Biochemistry and Physiology, Part B 138: 265-276.