Douglas Emlen

Douglas Emlen



Personal Website
Curriculum Vitae


BIOB 272 Genetics & Evolution  (Spring)

BIOE 406 & 409 Behavior & Evolution (alt Fall)




B.A. Cornell University, 1989
Ph.D. Princeton University, 1994

Research Interests

I am an evolutionary biologist and Professor of Biology at the University of Montana. My research provides insights into the development and evolution of exaggerated male weaponry, such as the horns found in scarab beetles. I combine approaches from behavioral ecology, genetics, phylogenetics, and developmental biology to understand how evolution has shaped these bizarre structures. Current projects include an examination of how altered expression of appendage patterning genes contributes to species differences in the shape of horns, and how the insulin receptor (InR) pathway modulates the size of male weapons in response to the larval nutritional environment.  I also communicate the excitement of evolutionary biology to the popular press, such as National Public Radio and the New York Times, contributing to public understanding of animal diversity and morphological evolution.


MeatEater Podcast with Steve Rinella, Aug 2019:

Full Length Documentary for NOVA: Extreme Animal Weapons (now available on Netflix):

Sci Show Interview with Hank Green, number 1: Feb 2015 (

Sci SHow Interview with Hank Green, number 2: August 2015 (

Science For the People Interview: August 2015 (

NPR On Point Interview November 2014 (

NPR Science Friday Interview November 2014 (


Selected Publications


2019. O'Brien, Devin M., Boisseau, Romain P., Duell, Meghan, McCullough, Erin, Powell, Erin C., Somjee, Ummat, Solie, Sarah, Hickey, Anthony J., Holwell, Gregory L., Painting, Christina J., Emlen, Douglas J. Muscle mass drives cost in sexually selected arthropod weapons. Proceedings of the Royal Society of London, Series B 286: 20191063.

2018.  Metz, M.C., Emlen, D.J., Stahler, D.R., MacNulty, D.R., Smith, D.W., Hebblewhite, M. Predation shapes the evolutionary traits of cervid weapons. Nature Ecology & Evolution 2: 1619–1625.

2018. Ohde, T., Morit, S., Shigenobu, S., Morita, J., Mizutani, T., Gotoh, H.,     Zinna, R. A., Nakata, M., Ito, Y., Wada, K., Kitano, Y., Yuzaki, K., Toga,     K., Mase, M., Kadota, K., Rushe, J., Corley Lavine, L., Emlen, D. J., Niimi,T. Rhinoceros beetle horn development reveals deep parallels with dung beetles. PLoS Genetics 14: e1007651.

2018. O'Brien, D. M., Allen, C. E., Van Kleeck. M. J., Hone, D., Knell, R., Knapp, A., Christiansen S., Emlen, D. J. On the evolution of extreme structures: static scaling and the function of sexually selected signals. Animal Behaviour 144: 95-108

2018. Zinna. R., Emlen, D., Lavine, L. C., Johns, A., Gotoh, H., Niimi, T., Dworkin, I. Sexual dimorphism and heightened conditional expression in a sexually selected weapon in the Asian rhinoceros beetle. Molecular Ecology 27: 5049-5072.

2017.  O’Brien, D. M., Katsuki, M., Emlen D. J. Selection on an extreme weapon in the frog legged leaf beetle (Sagra femorata). Evolution. DOI: 10.1111/evo.13336. (6)

2017. Gotoh, H., Zinna, R. A., Ishikawa, Y., Miyakawa, H., Ishikawa, A., Sugime, Y., Emlen, D. J., Lavine, L. C., Miura, T. The function of appendage patterning genes in mandible development of the sexually dimorphic stag beetle. Developmental Biology 422: 24-32. 

2016. McCullough, E., Miller, C. W., Emlen, D. J. Why sexually selected weapons are not ornaments. Trends in Ecology & Evolution 31: 742-751. (With Cover) (38)

2016. Cooley, N.L., Emlen, D. J., Woods, H. A. Self-heating by largfe insect larvae? Journal of Thermal Biology 62: 76-83.

2016. Gotoh, H., Zinna, R., Warren, I., DeNieu, M., Niimi, T., Dworkin, I., Emlen, D., Miura, T., Lavine, L. Identification and functional analyses of sex determination genes in the sexually dimorphic stag beetle Cyclommatus metallifer. BMC Genomics, 17: 250. 

2015. Lavine, L., Gotoh, H., Brent, C. S., Dworkin, I. and Emlen, D. 2014. Exaggerated trait growth in insects. Annual Review of Entomology, 60:453-72.

2014. McCullough, E., Tobalske, B., and Emlen, D. J. Structural adaptations to diverse fighting styles in sexually selected weapons. Proceedings of the National Academy of Sciences. doi: 10.1073/pnas.1409585111.

2014. Gotoh, H. Miyakawa, H., Ishikawa, A., Ishikawa, Y., Sugime, Y., Emlen, D. J., Lavine, L. C., and Miura, T. Developmental link between sex and nutrition: doublesex regulates sex-specific mandible growth via juvenile hormone signaling in stag beetles. PLoS Genetics, journal.pgen.1004098. (With Cover)

2013. Warren, I. A., Gotoh, H., Dworkin, I. M., Emlen, D. J. and Corley-Lavine, L. A. A general mechanism for conditional expression of exaggerated sexually- selected traits. BioEssays, 35: 889-899. 

2012. Â  Emlen, D.J., Warren, I. A., Johns, A., Dworkin, I. and Corley-Lavine, L. A     mechanism of extreme growth and reliable signaling in sexually selected     ornaments and weapons. Science, 337: 860-864. (With Cover) 




2019 BEETLE BATTLES: One Scientist's Journey of Adventure and Discovery (Roaring Brook/Macmillan)

- A "narrative nonfiction" book about science for middle school readers


2019 EVOLUTION: Making Sense of Life, 3rd Edition (Macmillan Learning)


2014 ANIMAL WEAPONS: The Evolution of Battle (Henry Holt/ Macmillan Publishers)

Winner: Phi Beta Kappan Science Book of the Year (2015)



Elected to the American Academy of Arts and Sciences (2016).

Phi Beta Kappa Science Book of the Year Award (2015), Animal Weapons: The Evolution of Battle (Henry Holt).

Carnegie/CASE Professor of the Year Award for the state of Montana (2015).

Distinguished Teaching Award, The University of Montana (2014).

E. O. Wilson Naturalist Award, The American Society of Naturalists (2013).

Presidential Early Career Award, Office of Science and Technology Policy, Executive Office of the President, Washington, D.C. (2002)

American Naturalists Young Investigator Prize (1997).

Neuroscience Courses

BIOE 406 - Behavior & Evolution

Home Department

Division of Biological Sciences 

Area of Expertise

Animal Weaponry; Beetles; Evolutionary Biology