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in review

Schwab DB, Casasa A, Moczek AP (in review). Developmental niche construction differentially affects growth, scaling, and sexual dimorphism in three species of dung beetles.

Macagno ALM, Zattara EE, Ezeakudo O, Moczek AP, Ledon-Rettig CC (in review). Adaptive maternal behavioral plasticity and developmental programming mitigate the transgenerational effects of temperature in dung beetles.

Parzer H, Polly D, Moczek AP (in revision). The evolution of size and shape: insights from the genitalia of dung beetles.

in press / 2017

Moczek AP 2017. The shape of things to come: evo devo perspectives on causes and consequences in evolution. In: Cause and Process in Evolution; edited by K. Laland and T. Uller. Vienna Series in Theoretical Biology, in press.

Schwab DB, Moczek AP (2017). Evo devo and niche construction. In: Evolutionary Developmental Biology – A Reference Guide, edited by Laura Nuño de la Rosa and Gerd B. Müller. Springer, in press.

Beckers O, Kijimoto T, Moczek AP 2017. doublesex alters aggressiveness as a function of social context and sex in the polyphenic beetle Onthophagus taurus. Animal Behavior, in press.

Pespeni MH, Ladner JT, Moczek AP 2017. Signals of selection in conditionally expressed genes in the diversification of three horned beetles species. Journal of Evolutionary Biology [pdf]

Casasa S, Schwab DB, Moczek AP 2017. Developmental regulation and evolution of scaling: novel insights through the study of Onthophagus beetles.  Current Opinion in Insect Science 19: 52–60 [pdf]

Ledon-Rettig CC, Zattara EE, Moczek AP 2016. Asymmetric interactions between doublesex and sex- and tissue-specific target genes mediate sexual dimorphism in beetles. Nature Communications 8:14593 [pdf]


Schwab DB, Moczek AP 2016. Nutrient stress during ontogeny alters patterns of resource allocation in two species of horned beetles. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology 325:481-490. [pdf]

McKenna DD, Scully ED, Pauchet Y….Moczek AP (+65 co-autors) 2016. Genome of the Asian longhorned beetle (Anoplophora glabripennis), a globally significant invasive species, reveals key functional and evolutionary innovations at the beetle-plant interface. Genome Biology 17:227. [pdf]

Schwab DB, Riggs HE, Newton ILG, Moczek AP 2016. Developmental and ecological benefits of the maternally transmitted microbiota in a dung beetle. American Naturalist 188: 679-692. [pdf]

Zattara EE, Busey H, Linz D, Tomoyasu Y, Moczek AP 2016. Neofunctionalization of embryonic head patterning genes facilitates the positioning of novel traits on the dorsal head of adult beetles. Proceedings of the Royal Society of London, Series B. 283: 20160824. [pdf]

Casasa S, Moczek AP 2016. -BOOK REVIEW- Ecological Developmental Biology: the Environmental Regulation of Development, Health, and Evolution, 2nd edition, by Scott F. Gilbert and David Epel, Sinauer. Quarterly Review of Biology 91: 513-514. [pdf]

Busey HA, Zattara EE, Moczek AP 2016. Conservation, innovation, and bias: embryonic segment boundaries position posterior, but not anterior, head horns in adult beetles. Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, 326: 271-279. [pdf]

Silva DP, Vilela B, Buzatto BA, Moczek AP, Hortal J 2016. Contextualized niche shifts upon independent invasions by the dung beetle Onthophagus taurusBiological Invasions 18: 3137-3148.. [pdf]

Kijimoto T, Moczek AP 2016. Hedgehog signaling enables nutrition-responsive inhibition of an alternative morph in a polyphenic beetle. Proceedings of the National Academy of Sciences, 113: 5982-5987. [pdf]

Ledon-Rettig CC, Moczek AP 2016. The transcriptomic basis of tissue- and nutrition-dependent sexual dimorphism in the beetle Onthophagus taurusEcology and Evolution 6(6): 1601–1613. [pdf]

Macagno ALM, Moczek AP, Pizzo A 2016. Rapid divergence of nesting depth and digging appendages among tunneling dung beetle populations and species. American Naturalist 187: E143-E151. [pdf]

Snell-Rood EC, Burger M, Hutton Q, Moczek AP 2016. Effects of parental care on the accumulation and release of cryptic genetic variation: review of mechanisms and a case study of dung beetles.  Evolutionary Ecology 30:251–265. [pdf]


Macagno ALM, Moczek AP 2015. Appendage-patterning genes regulate male and female copulatory structures in horned beetles. Evolution & Development 17: 248-245. [pdf]

Macagno ALM, Beckers M, Moczek AP (2015). Differentiation of early ovarian development and the evolution of fecundity in rapidly diverging exotic beetle populations. Journal of Experimental Zoology Part A 323: 679-688. [pdf]

Laland KN, Uller T, Feldman M, Sterelny K, Müller GB, Moczek AP, Jablonka E, Odling-Smee J 2015.  Darwin Review;  The extended evolutionary synthesis: its structure, assumptions, and predictions. Proceedings of the Royal Society of London, Series B, 282: 20151019. [pdf]

Moczek AP, Sears, KE, Stollewerk A, Wittkopp PJ, Diggle P, Dworkin I, Ledon-Rettig C, Matus DQ, Roth S, Abouheif E, Brown FD, Chiu C, Cohen CS, De Tomaso AW, Gilbert SF, Hall B, Love A, Lyons DC, Sanger T, Smith J, Specht C, Vallejo-Marin M, Extavour CG 2015. The significance and scope of evolutionary developmental biology: a vision for the 21st century. Evolution & Development, 17: 198–219. [pdf]

Moczek AP 2015. Developmental plasticity and evolution – quo vadis? Heredity 115, 302-305. [pdf]

Beckers OM, Anderson W, Moczek AP 2015. A combination of developmental plasticity, parental effects, and genetic differentiation mediates divergences in life history traits between dung beetle populations. Evolution & Development, 17: 148–159. [pdf]

Moczek AP 2015. Re-evaluating the environment in developmental evolution. Frontiers in Ecology and Evolution, 3:7. [pdf]

Moczek AP and Ledón Rettig C 2015. -BOOK REVIEW- Advances in Evolutionary Developmental Biology, edited by J. Todd Streelman; Wiley-Blackwell. Quarterly Review of Biology 90: 340-341.


KijimotoT, Snell-Rood EC, Rocha G, Pespeni M, Kafadar K Moczek AP 2014. The nutritionally responsive transcriptome of the polyphenic beetle Onthophagus taurus and the importance of sexual dimorphism and body region. Proceedings of the Royal Society of London, Series B 281: 20142084. [pdf]

Laland K, Tobias U, Feldman M, Sterelny K, Müller G, Moczek AP, Jablonka E, Odling-Smee J. 2014. Does evolutionary theory need a rethink? Nature 514: 161-164. [pdf]

Schwab DB, Moczek AP 2014. Resource allocation during ontogeny is influenced by genetic, developmental, and ecological factors in the horned beetle Onthophagus taurusProceedings of the Royal Society of London, Series B, in press. [pdf]

Stansbury M, Moczek AP 2014. The function of Hox and appendage patterning genes in the development of a novel organ, the Photuris firefly lantern. Proceedings of the Royal Society of London, Series B 281: 1471-2954. [pdf]

Moczek AP, Kijimoto T 2014. Development and evolution of insect polyphenisms: novel insights through the study of sex determination mechanisms. Current Opinion in Insect Science, 1: 52-58. [pdf]

Moczek AP, KijimotoT, Snell-Rood EC, Rocha G, Pespeni M, Kafadar K 2014. Evolutionary and ecological genomics of plasticity: novel approaches and first insights from the study of horned beetles. In: Ecological Genomics; edited by C. Landry and N. Aubin-Horth. Springer Verlag, Berlin. pp. 127-148. [pdf]

Moczek AP 2014. Towards a theory of development through a theory of developmental evolution. In: Towards a  Theory of Development; edited by A. Minelli & T. Pradeu, Oxford University Press. [pdf]


Estes AM, Hearn DJ, Snell-Rood EC, Feindler M, Feeser K, Abebe T, Dunning JC Hotopp, Moczek AP 2013. Brood ball-mediated transmission of microbiome members in the dung beetle, Onthophagus taurus (Coleoptera: Scarabaeidae). PLoS ONE 8(11): e79061.

Stansbury M, Moczek AP 2013. The evolvability of arthropods. In: Arthropod Biology and Evolution: Molecules, Development, Morphology; edited by A. Minelli, G Boxshall and G Fusco. Springer Verlag, Berlin, pp. 479-493. [pdf]

Snell-Rood EC, Moczek AP 2013. Horns and the role of development in the evolution of beetle contests. In: Animal Contests. Ed. by: ICW Hardy & M Briffa. Cambridge University Press, UK. pp. 178-198.

Snell-Rood EC, Troth A, Moczek AP 2013. DNA Methylation as a mechanism of nutritional plasticity: insights from horned beetles. Journal of Experimental Zoology 320: 22–34. [pdf]


Kijimoto T, Moczek AP, Andrews J 2012. Diversification of doublesex function regulates morph-, sex-, and species-specific expression of beetle horns. Proceedings of the National Academy of Sciences. 10.1073/pnas.1118589109 [pdf]

Moczek AP 2012. – BOOK REVIEW – Endless brains most beautiful: A review of Arthropod Brains: Evolution, Functional Elegance and Historical Significance, by Nicholas Strausfeld. Harvard University Press. Evolution & Development 14: 537–538. [pdf]

Kijimoto T, Pespeni M, Beckers O, Moczek AP 2012. Beetle horns and horned beetles: emerging models in developmental evolution and ecology. WIREs Interdisciplinary Reviews in Developmental Biology. 10.1002/wdev.81. [pdf]

Snell-Rood EC, Moczek AP 2012. Insulin signaling as a mechanism underlying developmental plasticity: the role of FOXO in a nutritional polyphenism. PLoS ONE. 7(4): e34857. [pdf]

Moczek AP 2012. The nature of nurture and the future of evodevo: toward a comprehensive theory of developmental evolution. Integrative and Comparative Biology. 52: 108-119. [pdf]

Valena S, Moczek AP 2012. Epigenetic mechanisms underlying developmental plasticity in horned beetles. In: The Epigenetics of Emerging and Nonmodel Organisms. Genetics Research International. 2012: 576303. [pdf]

Moczek AP, Valena S 2012. – BOOK REVIEW – Epigenetics: Linking Genotype and Phenotype in Development and Evolution, edited by Benedikt Hallgrßmsson and Brian K. Hall. University of California Press. Quarterly Review of Biology. 87: 69-70.

Wasik B, Moczek AP 2012. pangolin expression influences the development of a morphological novelty: beetle horns. Genesis. 50: 404-414. [pdf]


Macagno, AM, Pizzo A, Palestrini C, Rolando A, Moczek AP 2011. Codivergence between male and female copulatory structures in Onthophagus beetles. PLoS ONE. 6(12): e28893. [pdf]

Moczek AP, Sultan S, Foster S, Ledon-Rettig C, Dworkin I, Nijhout HF, Abouheif E, Pfennig D (2011). The role of developmental plasticity in evolutionary innovation. Proceedings of the Royal Society of London, Series B. 278: 2705-2713. [pdf]

Moczek AP 2011. The origins of novelty. Nature 473: 34-35. (invited commentary) [pdf]

Wasik B, Moczek AP 2011. decapentaplegic (dpp) regulates the growth of a morphological novelty, beetle horns. Development, Genes and Evolution, 221: 17-27. [pdf]

Simonnet F, Moczek AP 2011. Conservation and diversification of gene function during mouthpart development in Onthophagus beetles. Evolution & Development, 13: 280-289. [pdf]

Moczek AP. 2011. Evolution and development: Onthophagus beetles and the evolutionary developmental genetics of innovation, allometry, and plasticity. In: Dung Beetle Ecology and Evolution, edited by Leigh W. Simmons & James Ridsdill-Smith; Wiley-Blackwell. p 126-151. [pdf]

Snell-Rood, E. C., A. Cash, M. V. Han, T. Kijimoto, A. Andrews, and A. P. Moczek 2011. Developmental decoupling of alternative phenotypes: insights from the transcriptomes of horn-polyphenic beetles. Evolution, 65: 231-245. [pdf]


Choi J-H, Kijimoto T, Snell-Rood EC, Tae H, Yang Y-I, Moczek AP, Andrews J. 2010. Gene discovery in the horned beetle Onthophagus taurus. BMC Genomics 11:703. [pdf]

Kijimoto T, Andrews J, Moczek AP 2010. Programmed cell death shapes the expression of horns within and between species of horned beetles. Evolution & Development, 12: 449-458. [pdf]

Pfennig D, Wund MA, Snell-Rood EC, Cruickshank T, Schlichting CD, Moczek AP 2010. Phenotypic plasticity’s impacts on diversification and speciation. Trends in Ecology and Evolution 25: 459-467. [pdf]

Wasik BR, Rose DJ, Moczek AP 2010. Beetle horns are regulated by the Hox gene, Sex combs reduced, in a species- and sex-specific manner. Evolution & Development 12: 353-362. [pdf]

Snell-Rood EC, VanDyken JD, Cruickshank TE, Wade MJ, Moczek AP 2010. Toward a population genetic framework of developmental evolution: costs, limits, and consequences of phenotypic plasticity. Bioessays 32: 71-81. [pdf]

Moczek AP 2010. Phenotypic plasticity and diversity in insects. In From polyphenism to complex metazoan life cycles, edited by A. Minelli, G Fusco Philosophical Transactions of the Royal Society of London, Series B 365: 593-603. [pdf]

Moczek AP 2010. – BOOK REVIEW – In the Light of Evolution, vol. 1: Adaptation and Complex Design, edited by John C. Avise and Francisco Ayala. National Center for Science Education 30:1-2


Kijimoto T, Costello J, Tang Z, Moczek AP, Andrews J 2009. EST and microarray analysis of horn development in Onthophagus beetles. BMC Genomics 2009, 10:504. [pdf]

Moczek AP 2009 . On the origins of novelty and diversity in development and evolution: a case study on beetle horns.In Cold Spring Harbor Annual Symposium Volume 74, Evolution: The Molecular Landscape. Cold Spring Harbor Laboratory Press.

Moczek AP, Rose, DJ 2009. Differential recruitment of limb patterning genes during development and diversification of beetle horns. Proceedings of the National Academy of Sciences 106: 8992-8997.(Faculty of 1000 reviews)[pdf]

Moczek AP 2009. The origin and diversification of complex traits through micro- and macro-evolution of development: Insights from horned beetles. Current Topics in Developmental Biology (CTDB). Ed. by: W Jeffery. Elsevier / Academic Press, New York, vol. 86: 135-162. [pdf]

Tomkins J, Moczek AP 2009. Patterns of threshold evolution in polyphenic insects under different developmental models.Evolution 62: 459-468. [pdf]

Moczek AP 2009. Developmental plasticity and the origins of diversity: a case study on horned beetles. In: Phenotypic plasticity in insects: mechanisms and consequences. pp. 81-134. Edited by: TN Ananthakrishnan & D Whitman. Science Publishers, Inc. Plymouth, UK. [pdf]

Moczek AP – BOOK REVIEW – Endless forms most strange: A review of: The superorganism: the beauty, elegance, and strangeness of insect societies. Evolution and Development 11: 754-756. [pdf]


Parzer HF, Moczek AP 2008. Rapid antagonistic coevolution between primary and secondary sexual characters in horned beetles. Evolution 62-9: 2423-2428. [pdf]

Moczek AP, Snell-Rood EC 2008. The basis of bee-ing different: the role of gene silencing in plasticity (Commentary). Evolution and Development 10: 511-513. [pdf]

Moczek AP 2008. On the origin of novelty in development and evolution. Bioessays 5: 432-447. [pdf]

Shepherd BL*, Prange HD, Moczek AP 2008. Some like it hot: body and weapon size affect thermoregulation in horned beetles.Journal of Insect Physiology 54: 604-611. [pdf]

Pizzo A, Roggero A, Palestrini C, Moczek AP, Rolando A 2008. Rapid shape divergences between natural and introduced populations of a horned beetle partly mirror divergences between species. Evolution and Development 10: 166-175. [pdf]


Moczek AP 2007. Pupal remodeling and the evolution and development of alternative male morphologies in horned beetles.BMC Evolutionary Biology 7: 151. [pdf]

Moczek AP, Andrews J, Kijimoto T, Yerushalmi Y, Rose D 2007. Emerging model systems in evo-devo: horned beetles and the origins of diversity. Evolution and Development 9: 323-328. [pdf]

Moczek AP 2007. Developmental capacitance, genetic accommodation, and adaptive evolution. Evolution and Development 9: 299-305. [pdf]

Shelby JA*, Madewell R*, Moczek AP 2007. Juvenile hormone mediates sexual dimorphism in horned beetles. Journal of Experimental Zoology B, Molecular and Developmental Evolution 308B: 417-427. [pdf]


Moczek AP, Cruickshank, TE, Shelby JA* 2006. When ontogeny reveals what phylogeny hides: gain and loss of horns during development and evolution of horned beetles. Evolution 60: 2329-2341. [pdf]

Moczek AP 2006. Pupal remodeling and the development and evolution of sexual dimorphism in horned beetles. The American Naturalist 168: 711-729. (American Naturalist Feature article of the month, Faculty 1000 review) [pdf]

Moczek AP, Rose D, Sewell W, Kesselring BR 2006. Conservation, innovation, and the evolution of horned beetle diversity.Development Genes and Evolution 216: 655-665. [pdf]

Moczek AP 2006. Integrating micro- and macroevolution of development through the study of horned beetles.Heredity 97: 168-178. [pdf]

Madewell R*, Moczek AP 2006. Horn possession reduces maneuverability in a horn-polyphenic beetle. Journal of Insect Science 6.21. [pdf]

Moczek AP 2006. A matter of measurements: challenges and approaches in the comparative analysis of static allometries. The American Naturalist 167: 606-611. [pdf]

Moczek AP, *Cochrane J 2006. Intraspecific female brood parasitism in the dung beetle Onthophagus taurus. Ecological Entomology 31: 1-6. [pdf]

Moczek AP 2006. – BOOK REVIEW- The origins of diversity: A review of Evolution of the Insects, by David Grimaldi and Michael S. Engel. Evolution & Development 8: 111-112. [pdf]


Moczek AP 2005. The evolution and development of novel traits, or how beetles got their horns. BioScience 11: 935-951.(Faculty of 1000 review) [pdf]

Moczek AP, Nagy LM 2005. Diverse developmental mechanisms contribute to different levels of diversity in horned beetles.Evolution & Development 7: 175-185. [pdf]


Moczek AP, Bruehl, CB, Krell FTK 2004. Linear and threshold dependent expression of secondary sexual traits in the same individual: insights from a horned beetle. Biological Jounal of the Linnean Society 83: 473-480 [pdf]

Moczek AP, Nijhout HF 2004. Tradeoffs during the development of primary and secondary sexual traits in a dimorphic beetle.The American Naturalist 163: 184-191 [pdf]


Moczek AP 2003. The behavioral ecology of threshold evolution in a polyphenic beetle. Behavioral Ecology 14: 831-854 [pdf]

Moczek AP, Nijhout HF 2003. Rapid evolution of a polyphenic threshold. Evolution & Development 5: 259-268 [pdf]


Moczek AP 2002. Allometric plasticity in a polyphenic beetle. Ecological Entomology 27: 58-67 [pdf]

Moczek AP, Nijhout HF 2002a. Developmental mechanisms of threshold evolution in a polyphenic beetle. Evolution & Development 4: 252-264 [pdf]

Moczek AP, Nijhout HF 2002b. A method for sexing third instar larvae of the genus Onthophagus LATREILLE (Coleoptera: Scarabaeidae). Coleopterist Bulletin 56: 279-284 [pdf]

Moczek AP, Hunt J, Emlen DJ, Simmons LW 2002. Evolution of a developmental threshold in exotic populations of a polyphenic beetle. Evolutionary Ecology Research 4: 587-601 [pdf]

Moczek AP 2002. – BOOK REVIEW- How complexity pervades biology . Complexity 7: 16-17 [pdf]


*Shafiei M, Moczek AP, Nijhout HF 2001. Food availability controls onset of metamorphosis in the dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae). Physiological Entomology 26: 173-180 [pdf]


Moczek AP, Emlen DJ 2000. Male horn dimorphism in the scarab beetle Onthophagus taurus: do alternative reproductive tactics favor alternative phenotypes? Animal Behaviour 59: 459-466 [pdf]

Moczek AP 1999. Facultative paternal investment in the polyphenic beetle Onthophagus taurus: the role of male morphology and social context. Behavioral Ecology 10: 641-647 [pdf]

Moczek AP, Emlen DJ 1999. Proximate determination of male horn dimorphism in the beetle Onthophagus taurus(Coleoptera: Scarabaeidae). Journal of Evolutionary Biology 12: 27-37 [pdf]

Moczek AP 1998. Horn polyphenism in the beetle Onthophagus taurus: diet quality and plasticity in parental investment determine adult body size and male horn morphology. Behavioral Ecology 9: 636-641 [pdf]