Cooley, N., Emlen, D.J., and Woods, H.A. (2016) Self-heating by large insect larvae? Journal of Thermal Biology 62: 76-83. [PDF]

Gotoh, H., Zinna, R.A., Warren, I., DeNieu, M., Niimi, T., Dworkin, I., Emlen, D.J., Miura, T., Lavine, L.C. (2016) Identification and functional analyses of sex determination genes in the sexually dimorphic stag beetle Cyclommatus metalliferBMC Genomics 17: 250-267. [PDF]

Lin, X., Yao, Y., Wang, B., Emlen, D.J., and Lavine, L.C. (2016) Ecological trade-offs between migration and reproduction are mediated by the nutrition-sensitive insulin pathway. International Journal of Biological Sciences 12: 607-616. [PDF]

McCullough, E.L., Miller, C.W., and Emlen, D.J. (2016) Why sexually selected weapons are not ornaments. Trends in Ecology and Evolution 31: 742-751. [PDF]

Smith, J.N., Emlen, D.J., and Pearson, D.E. (2016) Linking native and invader traits explains native spider responses to plant invasions. PLOS ONE 11(4): e0153661. [PDF]


Gotoh, H., Hust, J.A., Miura, T., Niimi, T., Emlen, D.J., and Lavine, L.C. (2015) The fat/hippo signaling pathway links within-disc morphogen patterning to whole-animal signals during phenotypically plastic growth in insects. Developmental Dynamics 244: 1039-1045.[PDF]

Lavine, L., Gotoh, H., Brent, C., Dworkin, I., and Emlen, D.J. (2015) Exaggerated trait growth in insects. Annual Review of Entomology 60: 453-472.[PDF]

McCullough, E.L., Ledger, K.J., and Moore, T.Y. (2015) Variation in cross-sectional horn shape within and among rhinoceros beetle species. Biological Journal of the Linnean Society 115(4): 810-817.[PDF]

McCullough, E.L., Ledger, K.J., O’Brien, D.M., and Emlen, D.J. (2015) Variation in the allometry of exaggerated rhinoceros beetle horns. Animal Behaviour 109: 133-140.[PDF]

Gotoh, H., Miyakawa, H., Ishikawa, Y., Sugime, Y., Emlen, D.J., Lavine, L.C., and Miura, T. (2014) Developmental link between sex and nutrition; doublesex regulates sex-specific mandible growth via juvenile hormone signaling in stag beetles. PLoS Genetics 10(1): e1004098.[PDF]

Johns, A., H. Gotoh, E. L. McCullough, D. J. Emlen, and L. C. Lavine (2014) Heightened condition-dependent growth of sexually selected weapons in the rhinoceros beetle, Trypoxylus dichotomous (Coleoptera: Scarabaeidae). Integrative and Comparative Biology, doi:10.1093/icb/icu041. [PDF]

McCullough, E.L.(2014) Mechanical limits to maximum weapon size in a giant rhinoceros beetle. Proceedings of the Royal Society. B. Biological Sciences 281: 20140696, doi:10.1098/rspb.2014.0696. [PDF]

McCullough, E.L., Tobalske, B.W., Emlen, D.J. (2014) Structural adaptations to diverse fighting styles in sexually selected weapons. Proceedings of the National Academy of Sciences 111(40): 14484-14488, doi:10.1073/pnas.1409585111. [PDF]

Warren, I. A., J. C. Vera, A. Johns, R. Zinna, J. H. Marden, D. J. Emlen, I. Dworkin, and L. C. Lavine (2014) Insights into the development and evolution of exaggerated traits using de novo transcriptomes of two species of horned scarab beetles. PLOS One 9(2): e88364, doi:10.1371/journal.pone.0088364. [PDF]

Hattori, A., Sugime, Y., Sasa, C., Miyakawa, H., Ishikawa, Y., Miyazaki, S., Okada, Y., Cornette, R., Corley-Lavine, L., Emlen, D.J., Koshikawa, S., and Miura, T. (2013) Soldier morphogenesis in the damp-wood termite is regulated by the insulin signaling pathway. Journal of Experimental Zoology (Molecular and Developmental Evolution) 320B: 295-306.[PDF]

Corley Lavine, L., Hahn, L.L., and Warren, I. (2013) Cloning and characterization of an mRNA encoding an insulin receptor from the horned scarab beetle Onthophagus nigriventris(Coleoptera:Scarabaeidae). Archives of Insect Biochemistry and Physiology 82:43-57.[PDF]

McCullough, E.L. (2013) Using radio telemetry to assess movement patterns in a giant rhinoceros beetle: are there differences among majors, minors, and females? Journal of Insect Behavior 26:51-56.[PDF]

McCullough, E.L. and Emlen, D.J. (2013) Evaluating the costs of a sexually selected weapon: big horns at a small price. Animal Behaviour 86:977-985.[PDF]

McCullough, E.L. and Tobalske, B. (2013) Elaborate horns in a giant rhinoceros beetle incur negligible aerodynamic costs. Proceedings of the Royal Society. B. Biological Sciences 280:20130197.[PDF]

McCullough, E.L. and Zinna, R. (2013) Sensilla density corresponds to thermions of the horn most frequently used during combat in the giant rhinoceros beetle Trypoxylus dichotomus (Coleoptera: Scarabaeidae: Dynastinae). Annals of The Entomological Society of America 106(4): 518-523.[PDF]

Warren, I.A., Gotoh, H., Dworkin, I.M., Emlen, D.J., and Lavine, L.C.(2013) A general mechanism for conditional expression of exaggerated sexually-selected traits. Bioessays 35:889-899.[PDF]

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

McCullough, E., Weingarden, P., and Emlen, D. J. (2012) Costs of elaborate weapons in a rhinoceros beetle:    how difficult is it to fly with a big horn? Behavioral Ecology, doi:10.1093/beheco/ars069. [PDF]

Van Truong, T., Byun, D., Corley-Lavine, L., Emlen, D.J., Park, H. C. and  Kim, M. J. (2012) Flight     behavior of the rhinoceros beetle Trypoxylus dichotomus during electrical nerve stimulation.  Bioinspiration & Biomimetics 7: 036021 (11pp). [PDF]

Hartfelder, K. and Emlen, D. J. (2011) Endocrine control of insect polyphenism. Insect Endocrinology, L. I. Gilbert, Ed., Elsevier. (Pp. 464-522) [PDF]

Gotoh, H., Cornette, R., Koshikawa, S., Okada, Y., Corley Lavine, L., Emlen, D. J., and Miura, T. (2011) Juvenile Hormone Regulates Extreme Mandible Growth in Male Stag Beetles. PloS One 6(6):e21139. [PDF]

Emlen, D. J. (2010) A developmental view of exaggerated growth and conditional expression in the weapons of sexual selection. In: P. R. Grant and B. R. Grant, eds. “From Field Observations to Mechanisms: A Program in Evolutionary Biology” Princeton University Press Pp. 149-171.

Emlen, D. J. (2010) Diversity in the weapons of sexual selection: Horn evolution in the beetle genus Onthophagus.  In: Evolution Emerging: Essays From the Field and Laboratory From Leading  Evolutionary Biologists. Ed: Jonathan Losos. Roberts & Company Publishers Pp. 149-170. [PDF]

Miller, C. W. and Emlen, D. J. (2009) Across and within population differences in the size and scaling relationship of a sexually-selected trait in Leptoscelis tricolor (Hemiptera: Coreidae). Annals  of the Entomological Society of America 103: 209-215.[PDF]

Miller, C. W. and Emlen, D. J. (2010) Dynamic effects of oviposition site on offspring sexually-selected traits and scaling relationships. Evolutionary Ecology 24: 375-390. [PDF]

Woods, H. A., and Smith, J.N. (2010) Universal model for water costs of gas exchange by animals and plants. Proceedings of the National Academy of Sciences of the United States of America 107:8469-8474. [PDF]

Rowland, J. M. and Emlen, D. J. (2009) Two thresholds, three male forms result in facultative male trimorphism in beetles. Science 323: 773-777. [PDF]

________  Highlighted in Nature (Feb. 2009); Scarabs (August 2009).

Woods, H. A., Sprague, J.C., and Smith, J.N. (2009) Cavitation in the embryonic tracheal system of Manduca sexta. Journal of Experimental Biology 212:3296-3304.[PDF]

Emlen, D. J. (2008) The evolution of animal weapons. Annual Review of Ecology, Systematics and Evolution. 39: 387-413. [PDF]

________ Highlighted in The New York Times (March 23, by Nicholas Wade);  Tages-Anzeiger (April 2009); El Mercurio (March 2009). 

Simmons, L. W. and Emlen, D. J.  (2008) No fecundity cost of female secondary sexual trait expression in the horned beetle Onthophagus sagittarius. Journal of Evolutionary Biology 21: 1227-  1235. (with Cover) [PDF]

Frankino, W. A., Emlen, D. J. and Shingleton, A. W. (2008) Experimental approaches to studying the evolution of animal form: The shape of things to Come. In: Experimental Evolution: Concepts, Methods, and Applications of Selection Experiments. T. Garland, Jr. and M. R. Rose, editors. University of California Press.

Emlen, D. J. The role of genes and the environment in the expression and evolution of animal alternative tactics. Pp. 85-108 In: Alternative Reproductive tactics: an Integrative Approach. R. Oliveira, M. Taborsky and J. Brockmann, eds., Cambridge University Press. (2008) 

 Simmons, L. W., Emlen, D. J., and Tomkins, J. (2007) Sperm competition games between sneaks and guards: A comparative analysis using dimorphic male beetles. Evolution 61: 2684-2692. (with Cover). [PDF]

Emlen, D. J., Corley Lavine, L., and Ewen-Campen, B.  (2007) On the origin and evolutionary diversification of beetle horns. Proceedings of the National Academy of Sciences 104 supplement1:  8661-8668. [PDF]

Shingleton, A. W., Frankino, W. A., Flatt, T., Nijhout, H. F. and Emlen D. J. (2007) Size and shape: The developmental regulation of static allometry in insects. BioEssays 29: 536-548. [PDF]

Simmons, L. W. and Emlen, D. J. (2006) Evolutionary trade-off between weapons and testes. Proceedings of the National Academy of Sciences 103: 16346-16351 (with Cover). [PDF]

________ Highlighted in National Geographic News (October 16, by John Roach);  New Scientist (October 16, by John Pickrell); Science News (October 17,  by Susan Milius); South Asia News (October 17,  2006); ABC News  Online (October 17, 2006 by Judy Skatsoon); ScienceNOW (October 24,  2006); India News Channel (October 25, 2006); SciTini (November 6,  2006 by Karen Wiens);  Newsweek (Poland, November 11, 2006); The  Telegraph (India, December 4, 2006 by Monojit DasGupta); Pour La  Science (December 2006).  

________  Faculty of 1000 rating of 6.0 “must read” by Gabriele Sorci.

Emlen, D. J. and Philips, T. K. (2006) Phylogenetic evidence for an association between tunneling behavior and the evolution of horns in dung beetles (Coleoptera: Scarabaeinae). Coleopterists  Society Monograph 5: 47-56. [PDF]

Emlen, D. J., Szafran, Q., Corley, L.W. and Dworkin, I. (2006) Insulin signaling and limb-patterning: Candidate pathways for the origin and evolutionary diversification of beetle horns. Heredity  97: 179-191. [PDF]

________  Facutly of 1000 rating of ‘Recommended’ by Russell Bonduriansky

 Emlen, D. J., Marangelo, J., Ball, B. and Cunningham, C. W.  (2005) Diversity in the weapons of sexual selection: Horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution 59: 1060-1084. (with Cover) [PDF]

________ Highlighted in The Loom (“An inordinate fondness for beetle horns” by  Carl Zimmer, June 2005) 

________  Faculty of 1000 rating of 6: “must read” by Edmund Brodie (June 2005)

Emlen, D. J., Hunt, J., and Simmons, L. W. (2005) Evolution of sexual dimorphism and male dimorphism in the expression of beetle horns: phylogenetic evidence for modularity, evolutionary lability, and constraint. American Naturalist 166: S42-S68.[PDF]

Hartfelder, K. and Emlen, D. J. (2005) Endocrine control of insect polyphenism. Comprehensive Molecular Insect Science 3: 652-702. 


Emlen, D. J. and Allen, C. E. (2004) Genotype to Phenotype: Physiological control of trait size and scaling in insects. Integrative and Comparative Biology 43: 617-634. [PDF]


Moczek, A. P., Hunt, J., Emlen, D. J. and Simmons, L. W. (2002) Threshold evolution in exotic populations of a polyphenic beetle.  Evolutionary Ecology Research  4: 587-601. 

Emlen, D. J. (2001) Costs and the diversification of exaggerated animal structures. Science 291: 1534-1536. [PDF]

_________ Highlighted in This Week in Science, by Phil Szuromi, 291: 1443, in a Science's Compass Perspectives article: A horn for an eye, by Paul Harvey and Charles Godfray, 291: 1505-1506; Picked up by Evolution of Dung beetle's horn explained, by Lisa Onaga, Feb. 23, 2001; ScienceNow: On the horns of a dilemma, by Menno Schilthuizen, Feb. 28, 2001; Science Magazine Online: Sizing up dung beetle evolution, by Wade Roush, Feb. 28, 2001; Study Works! Online: You can't have your horns and big eyes too, by Rachel Clark, Mar. 30, 2001; Geo Magazine: Auge gegen horn, May, 2001 p. 204.

Emlen, D. J. and Nijhout, H. F. (2001) Hormonal Control of male horn length dimorphism in the dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae): A second critical period ofsensitivity to juvenile hormone. Journal of Insect Physiology. 47: 1045-1054.  

Emlen, D. J. (2000) Integrating development with evolution: A case study with beetle horns. BioScience. 50: 403-418. (with Cover) [PDF]

Emlen, D. J. (2000) Dig it, and they will come: A female dung beetle in her tunnel is sure to attract suitors big and small. Natural History May Issue: 64-69. 

Emlen, D. J. and Nijhout, H. F. (2000) The development and evolution of exaggerated morphologies in insects.  Annual Review of Entomology. 45: 661-708. [PDF]

Moczek, A. P. and Emlen, D. J. (2000) Male horn dimorphism in the scarab beetle Onthophagus taurus: do alternative tactics favor alternative phenotypes? Animal Behaviour. 59: 459-466. 

_________ Picked up by BBC Wildlife (September 2000, p. 35).


Stern, D.L.S. and Emlen, D.J. (1999) The developmental basis for allometry in insects. Development 126: 1091-1101. (with Cover) 

Emlen, D.J, and Nijhout, H. F. (1999) Hormonal control of male horn length dimorphism in the horned beetle Onthophagus taurus.  Journal of Insect Physiology 45: 45-53. 

Moczek, A, and Emlen, D.J. (1999) Proximate determination of male horn dimorphism in the beetle Onthophagus taurus (Coleoptera: Scarabaeidae). Journal of Evolutionary Biology 12: 27-37.  


Nijhout, H.F and Emlen, D.J.  (1998) Competition among body parts in the development and evolution of insect morphology. Proceedings of the National Academy of Sciences 95: 3685- 3689. [PDF]

_________  Selected for press release by P.N.A.S. and picked up by Science News (April 1998, p. 231), The New York Times (Science Tuesday, May 5, 1998), Earth Magazine (August, 1998, p. 11), Discover Magazine (September, 1998, p. 18-19).


Emlen, D.J. (1997) Alternative reproductive tactics and male dimorphism in the horned beetle Onthophagus acuminatus. Behavioral Ecology and Sociobiology 41: 335-341.1996.[PDF]

Emlen, D.J.  (1996) Artificial selection on horn length-body size allometry in the horned beetle Onthophagus acuminatus. Evolution 50: 1219-1230. [PDF]


 Emlen, D.J. (1994) Environmental control of horn length dimorphism in the beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Proceedings of the Royal Society of London, Series B, 256: 131-136. [PDF]


Emlen, D.J. (1993) New method for observing underground and soil surface behaviors. Coleopterists Bulletin, 47: 243-244.


Emlen, D.J.  (1992) Observations of a high altitude population of the Sage stem-galling fly Eutreta diana (Diptera: Tephritidae) and its associated parasitoids. Journal of the Kansas Entomological Society, 65: 203-207.


Emlen, S.T., Demong, N. J. and Emlen, D.J. (1989) Experimental induction of infanticide in female Wattled Jacanas. Auk, 106: 1-7.