The Page Lab

Sensory and Cognitive Ecology - Smithsonian Tropical Research Institute - Panama















Page, R.A., Jones, P.L. In press. Overcoming sensory uncertainty: factors affecting foraging decisions in frog-eating bats. In: Perception and Cognition in Animal Communication (volume editors, M.A. Bee and C.T. Miller)in the book series Animal Signals and Communication (series editors: P.K. McGregor and V.M. Janik). Springer.

ter Hofstede, H.M., Voigt-Heucke, S.L., Lang, A., Römer, H., Page, R.A., Faure, P.A., Dechmann, D.K.N. In press. Revisiting adaptations of Neotropical katydids (Orthoptera: Tettigoniidae) to gleaning bat predation. Neotropical Biodiversity.

Flores, E.E., Page, R.A., Cisneros, I., Rodriguez, S. In press. Oophaga vicentei (Vicente´s Poison Frog). Color change. Herpetological Review.

Gomes, D. G. E., Page, R. A., Geipel, I., Taylor, R. C., Ryan, M. J., Halfwerk W. 2016. Bats perceptually weight prey cues across sensory systems when hunting in noise. Science. 353: 1277-1280. doi: 10.1126/science.aaf7934 [pdf].

Halfwerk, W., Guerra, M.A., Lea, A.M., Page, R.A., Ryan, M.J. 2016. Amplitude regulation in frogs reveals the ancestral state of the Lombard effect. Behavioral Ecology. 27: 669-676. doi:10.1093/beheco/arv204 [pdf].

Jones, P.L., Page, R.A., Ratcliffe, J.M. 2016. To scream or to listen? Prey detection and discrimination in animal-eating bats. In: Bat Bioacoustics (volume editors: B. Fenton and A. Grinnell; series editor: A. Popper). Springer. Springer. pp. 93-116 [pdf].

Stange, N., Page, R.A., Ryan, M.J., Taylor, R.C. 2016. Interactions between complex multisensory signal components result in unexpected mate choice responses. Animal Behaviour. DOI: 10.1016/j.anbehav.2016.07.005 [pdf].

Symes, L.B., Page, R.A., ter Hofstede, H.M. In press. Interspecific variation in call timing by Neotropical forest katydids. Behavioral Ecology and Sociobiology. DOI 10.1007/s00265-016-2157-4 [pdf].

Ramakers J.J.C, Dechmann D.K.N., Page R.A., O’Mara M.T. (2016) Frugivorous bats prefer information from novel social partners. Animal Behaviour. 116: 83-87 [pdf].

Ripperger, S., Josic, D., Hierold, M., Koelpin, A., Weigel, R., Hartmann, M., Page, R.A., Mayer, F. 2016. Automated proximity sensing in small vertebrates: design of miniaturized sensor nodes and first field tests in bats. Ecology and Evolution. 6: 2179–2189 [pdf].

Trillo, P.A., Bernal, X.E., Caldwell, M., Halfwerk, W., Owens, M., Page, R.A. 2016. Collateral damage or a shadow of safety? The effects of signaling neighbors on the risks of parasitism and predation. Proceedings of the Royal Society Series B. 283: 20160343. [pdf].

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



Bader, E., Jung, K., Kalko, E.K.V., Page, R.A., Rodriguez, R., Sattler, T. 2015. Mobility explains the response of aerial insectivorous bats to anthropogenic habitat change in the Neotropics. Biological Conservation: 186: 97–106 [pdf].

Bulbert, M.W., Page, R.A., Bernal, X.E. 2015. Danger comes from all fronts: predator-dependent escape tactics of túngara frogs. PLoS ONE: 10(4): e0120546. doi:10.1371/journal.pone.0120546 [pdf].

Falk, J.J., ter Hofstede, H.M., Jones, P.L., Dixon, M.M., Faure, P.A., Kalko, E.K.V., Page, R.A. 2015. Sensory-based niche partitioning in a multiple predator-multiple prey community. Proceedings of the Royal Society Series B. 282: 20150520. [pdf].

Fugère, V., O'Mara, M.T., Page, R.A. 2015. Perceptual bias does not explain preference for prey call adornment in the frog-eating bat. Behavioral Ecology and Sociobiology. DOI 10.1007/s00265-015-1949-2 [pdf].

Rhebergen, F., Taylor, R.C., Ryan, M.J., Page, R.A., Halfwerk, W. 2015. Multimodal cues improve prey localisation under complex environmental conditions. Proceedings of the Royal Society Series B. 282: 20151403. [pdf].

Stockmaier, S., Dechmann, K.N., Page, R.A., O’Mara, M.T. 2015. No fever and leukocytosis in response to a lipopolysaccaride challenge in an insectivorous bat. Biology Letters. 11: 20150576. [pdf].



Jones, P.L., Ryan, M.J., Page, R.A. 2014. Population and seasonal variation in response to prey calls by an eavesdropping bat. Behavioral Ecology and Sociobiology. 68: 605-615. DOI 10.1007/s00265-013-1675-6 [pdf].

Halfwerk, W., Dixon, M.M., Ottens, K., Taylor, R.C., Ryan, M.J., Page, R.A., Jones, P.L. 2014. Risks of multimodal signaling: bat predators attend to dynamic motion in frog sexual displays. Journal of Experimental Biology. 217: 3038-3044. DOI 10.1242/jeb.107482 [pdf].

O’Mara, M.T., Dechmann, D.D, Page, R.A. 2014. Frugivorous bats evaluate the quality of social information when choosing novel foods. Behavioral Ecology. 1–7. doi:10.1093/beheco/aru120 [pdf].

Halfwerk, W., Page, R.A., Taylor, R.C., Wilson, P.S., Ryan, M.J. 2014. Cross-modal comparisons of signal components allow for relative distance assessment. Current Biology. 24:1751-1755 [pdf].

Clarin, T.M.A., Borissov, I., Page, R.A., Ratcliffe, J.M., Siemers, B.M. 2014. Social learning within and across species: information transfer in mouse-eared bats. Canadian Journal of Zoology. 92: 129-139. DOI 10.1139/cjz-2013-0211 [pdf].

Page, R.A., Ryan, M.J., Bernal, X.E. 2014. Be loved, be prey, be eaten. In: Animal Behavior, vol 3. Case Studies: Integration and Application of Animal Behavior (ed., K. Yasukawa), New York: Praeger. pp. 123-154 [pdf].

Halfwerk, W., Jones, P.L., Taylor, R.C., Ryan, M.J., Page, R.A. 2014. Risky ripples allow bats and frogs to eavesdrop on a multisensory sexual display. Science 342: 413-416, DOI 10.1126/science.1244812 [pdf].



Jones, P.L., Ryan, M.J., Page, R.A. 2013. When to approach novel prey cues? Social learning strategies in frog-eating bats. Proceedings of the Royal Society B. 280:20132330, [pdf].

Surlykke, A., Jakobsen, L., Kalko, E.K.V., Page, R.A. 2013. Echolocation intensity and directionality of perching and flying fringe-lipped bats, Trachops cirrhosus (Phyllostomidae). Frontiers in Physiology. 4:143, doi: 10.3389/fphys. 2013.00143 [pdf].

Clarin, T.M.A., Ruczyński, I., Page, R.A., Siemers, B.M. 2013. Foraging ecology predicts learning performance in insectivorous bats. PLoS ONE: 8(6): e64823 [pdf].

Jones, P.L., Farris, H.E., Ryan, M.J., Page, R.A. 2013. Do frog-eating bats perceptually bind the complex components of frog calls? Journal of Comparative Physiology A. 199: 279-283 [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: 505–509 [pdf].

Page, R.A., von Merten, S. (equal contribution), Siemers, B.M. 2012. Associative memory or algorithmic search: a comparative study on learning strategies of bats and shrews. Animal Cognition. 15: 495-504 [pdf].



Akre, K.L., Farris, H.E., Lea, A.M., Page, R.A., Ryan, M.J. 2011. Signal perception in frogs and bats and the evolution of mating signals. Science 333: 751-752 [pdf]; supporting online materials [pdf]; perspective by Rowe and Healy [pdf].

Jones, P.L., Page, R.A., Hartbauer, M., Siemers, B.M. 2011. Behavioral evidence for eavesdropping on prey song in two Palearctic sibling bat species. Behavioral Ecology and Sociobiology. 65: 333-340. [pdf].



Siemers, B.M. and Page, R.A. 2009. Behavioral studies of bats in captivity: Methodology, training, and experimental design. In: Ecological and Behavioral Methods for the Study of Bats. 2nd edition (ed. T.H. Kunz & S. Parsons), Baltimore: Johns Hopkins University Press. pp. 373-392 [pdf].

Bernal, X.E., Page, R.A., Ryan, M.J., Argo, T.F., and Wilson, P.S. 2009. Acoustic radiation patterns of mating calls of the túngara frog (Physalaemus pustulosus): implications for multiple receivers. Journal of the Acoustical Society of America. 126: 2757-2767 [pdf].


2008 and earlier

Page, R.A. and M.J. Ryan. 2008. The effect of signal complexity on localization performance in bats that localize frog calls. Animal Behaviour 76: 761-769 [pdf].

Bernal, X.E., Page, R.A., Rand, A.S., and Ryan, M.J. 2007. Cues for eavesdroppers: do frog calls indicate prey density and quality? The American Naturalist 169: 409-415 [pdf].

Page, R.A. 2007. Prey-predator communication: for your sensors only. Dispatch for Current Biology 17: R965-R966 [pdf].

Page, R.A. and M.J. Ryan. 2006. Social transmission of novel foraging behavior in bats: frog calls and their referents. Current Biology, 16: 1201-1205 [pdf]; dispatch by Ratcliffe and Fenton 2006 [pdf].

Page, R.A. and X.E. Bernal. 2006. Túngara frogs. Quick guide for Current Biology 16: R979-980 [pdf].

Page, R.A. and Ryan, M.J. 2005. Flexibility in assessment of prey cues: frog-eating bats and frog calls. Proceedings of the Royal Society Series B 272: 841-847 [pdf].

Gabor, C.R. and Page, R.A. 2003. Female preference for large males in sailfin mollies, Poecilia latipinna: the importance of predation pressure and reproductive status. Acta Ethologica 6: 7-12 [pdf].