Purdue University

Bernal Lab



Vencl, F., Ottens, K., Dixon, M.M., Candler, S., Bernal, X.E. Estrada, C. & Page. R.A. 2016. Pyrazine emission by a tropical firefly: an example of chemical aposematism? Biotropica doi:10.1111/btp.12336.

Trillo, P.A., Bernal, X.E., Caldwell, M.S., Halfwerk, W.H., Wessel, M.O. & Page. R.A. 2016. Collateral damage or a shadow of safety?: The effects of signalling heterospecific neighbours on the risks of parasitism and predation. Proc Roy Soc B 283: 20160343.

Aihara, I., Priyanka de Silva & Bernal, X.E. 2016. Acoustic preference of frog-Biting midges (Corethrella spp) attacking túngara frogs in their natural habitat. Ethology 122: 105-113. PDF

Beaty, L., Emmering, Q. & Bernal, X.E. 2016. Mixed sex effects on the second-to-fourth digit ratio of túngara frogs and cane toads. The Anatomical Record 299:421-427. PDF

Bernal, X.E. & Pinto, C.M. 2016. Sexual differences in prevalence of a new species of trypanosome that infects túngara frogs. International Journal of Parasitology: Wildlife and Parasites 5:40-47 PDF

Baugh, A., Ryan, M.J., Bernal, X.E. & Bee, M. 2016. Túngara frogs do not experience the continuity illusion. Behavioral Neuroscience. 130:62-74. PDF


Bernal, X.E. & de Silva, P. 2015. Cues used in host-seeking behavior by frog-biting midges (Corethrella spp Coquillet). Journal of Vector Ecology 40: 122-128. PDF

Bulbert, M.W., Page, P.A. & Bernal, X.E. 2015. Danger comes from all fronts: predator-dependent escape tactics of túngara frogs. PLoS One 10(4): e0120546. PDF

Beaty, L., Nyarko, K. & Bernal, X.E. 2015. Does light reveal their true toxicity? Light-induced changes in coloration through ontogeny in cane toad tadpoles (Rhinella marina). Herpetological Journal 25: 191-195. PDF

de Silva, P., Nutter, B. & Bernal, X.E. 2015. Use of acoustic mating signals in an eavesdropping frog-biting midge. Animal Behaviour 103:45-51. PDF

de Silva, P., Miranda, R. & Bernal, X.E. 2015. First report of mite parasitization in frog-biting midges (Corethrella species). International Journal of Parasitology 41:389-392 PDF

Candler, S.A. & Bernal, X.E. 2015. Differences in neophobia between cane toads from introduced and native populations. Behavior Ecology 26:97-104 PDF


Page, R.A., Ryan, M.J. & Bernal, X.E. 2014. Be Loved, Be Preyed, Be Eaten. In: Animal Behavior: Vols. 3. Yasakawa, K. (ed). ABC-CLIO Editorial. Santa Barbara, CA.

Akre, K., Bernal, X.E., & Ryan, M.J. 2014. Harmonic calls and indifferent females: Evidence against anuran origins for human consonance perception. Proceedings of the Royal Society B 281: 20140986 PDF

de Silva, P., Jaramillo C. & Bernal X.E. 2014. Selection of biting sites on anuran hosts by Corethrella Coquillett species. J Insect Behav 27:302-316 PDF


de Silva, P. C. & Bernal X.E. 2013. First report of the mating behavior of a species of frog-biting midge (Diptera: Corethrellidae). Florida Entomologist 96(4):1522-1529 PDF


Lloyd-Strovas, J. & Bernal, X.E. 2012. A review of undergraduate evolution education in United States Universities: Building a unifying framework. Evolution: Education and Outreach 5:453-46 PDF

Page, R.A., Schnelle, T., Kalko, E.K.V., Bunge, T. & Bernal, X.E. 2012. Reassessment of prey through sequential use of multiple sensory cues by an eavesdropping bat. Naturwissenschaften 99:505-50 PDF


Ryan, M.J., Bernal, X.E. & Rand, A.S. 2010. Female mate choice and the potential for ornament evolution in túngara frogs, Physalaemus pustulosus. Current Zoology 56:343-357 PDF


Bernal, X.E., Page, R.A., Ryan, M.J., Argo IV, T.F. & Wilson, P.S. 2009. Acoustic radiation patterns of mating calls of the túngara frog (Physalaemus pustulosus): Implications for multiple receivers. J Acous Soc Am 126:2757-276 PDF

Bernal, X.E., Akre, K.L., Baugh, A.T., Rand, A.S. & Ryan, M.J. 2009. Female and male behavioral response to advertisement calls of variable complexity in túngara frogs, Physalaemus pustulosus. Behav Ecol Sociobiol 63:1269-127 PDF

Bernal, X.E., Rand, A.S. & Ryan, M.J. 2009. Task differences underlie sexual dimorphism in behavior. Proc Roy Soc London B 276:1323-132 PDF


Cummings, M.E., Bernal, X.E., Reynaga, R., Rand, A.S. & Ryan M.J. 2008. Visual Sensitivity to a Conspicuous Male Cue Varies by Reproductive State in Physalaemus pustulosus females. J Exp Biol 211:1203-121 PDF


Ryan, M.J., Bernal, X.E., & Rand, S. 2007. Patterns of mating call preferences in túngara frogs, Physalaemus pustulosus. J Evol Biol 20:2235-224 PDF

Bernal, X.E., Rand, A.S. & Ryan, M.J. 2007. Sexual differences in the behavioral response of túngara frogs, Physalaemus pustulosus, to cues associated with increased predation risk. Ethology 113:755-763 PDF

Lampert, K.P., Bernal, X.E., Rand, A.S., Mueller, U.G. & Ryan, M.J. 2007. Island populations of Physalaemus pustulosus: history influences genetic diversity and morphology. Herpetologica 63(3):311-32 PDF

Bernal, X.E., Rand, A.S. & Ryan, M.J. 2007. Sexual differences in receiver permissiveness to advertisement calls in túngara frogs, Physalaemus pustulosus. Anim Behav 73:955-96 PDF

Bernal, X.E., Page, R.A., Rand, A.S. & Ryan, M.J. 2007. Cues for eavesdroppers: Do frog calls indicate prey density and quality?. Am Nat 169:412-41 PDF


Page, R.A & Bernal, X.E. 2006. Quick guide Túngara frogs. Current Biology 16(23):R979-R78 PDF

Bernal, X.E., Rand, A.S. & Ryan, M.J. 2006. Acoustic preferences and localization performance of blood-sucking flies (Corethrella Coquillett). Behav Ecol 17:709-71 PDF

Pauly, G.B., Bernal, X.E., Ryan, M.J. & Rand, A.S. 2006. The role of the vocal sac in increasing call rate in Physalaemus pustulosus. Physiol Biochem Zool 79:708-71 PDF

Lampert, K.P., Bernal, X.E., Rand A.S., Mueller, U.G. & Ryan, M.J. 2006. No evidence for female mate choice based on genetic similarity in the túngara frog, Physalaemus pustulosus. Behav Ecol Soc 59:796-80 PDF


Bernal, X.E, Guarnizo, C. and Luddecke, H. 2005. Geographic variation in the advertisement call and genetic structure of the Andean frog Colostethus palmatus. Herpetologica 61(4):395-40 PDF


Lüddecke, H., Amézquita, A., Bernal, X., Guzmán, F. 2000. Partitioning of vocal activity in a neotropical highland-frog community. Stud Neotrop Fauna & Environm 35:185-19 PDF


Bernal X., Guzmán F. 1999. The advertisement calls of three leutherodactylus species (Anura: Leptodactylidae) in a Colombian highland community. Rev Acad Colomb Cien 23 (Suplemento especial) 261-264 PDF


Lüddecke, H., Amézquita, A., Guzmán, F., Bernal, X., Agudelo, L. 1998. Comparison of environmental conditions and their effects on call characteristics of five frog species in a Colombian highland community. Miaud, C. & Guyétant, R. (eds). Le Bourget du Lac SEH:285-294 PDF

Natural History Notes

McCoy, K. & Bernal, X.E. 2013. Leptodactylus savagei (South American Bullfrog), Locomotion and Foraging. Natural History Note, Herp Review 44(3):496

Bernal, X.E. 2012. Leptodactylus fragilis (White-lipped Foamfrog), Distress Call. Natural History Note, Herp Review 43(2):32 PDF

Bernal, X.E. 2006. Caiman crocodilus (Spectacle caiman), Predation / Eavesdropper. Natural History Note, Herp Review 37(4):460-46 PDF

Pauly, G.B., Bernal, X.E. & Taylor, R. 2005. Scinax ruber (Red Snouted Treefrog), Arboreality and parachuting. Natural History Note, Herp Review 36(3):308-30 PDF

Bernal, X. & Ron, S. 2004. Leptodactylus fragilis (South American white-lipped frog), Courtship. Natural History Note, Herp Review 35(4):372-37 PDF