Publications of MJ Ryan

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2024/in press

Bredeson, J.V. et al. [MJR 1 of 35 authors]. 2024. Conserved chromatin and repetitive patterns reveal slow genome evolution in frogs. Nature Communications 15:579

Larter, L.C.; Ryan, M.J. 2024. Female preferences for more elaborate signals are an emergent outcome of male chorusing interactions in túngara frogs. The American Naturalist 203:92-108.

Larter,  L.C.; Ryan, M.J. 2024. Túngara frog call-timing decisions arise as internal rhythms interact with fluctuating chorus noise. Behavioral Ecology. 35(4), arae034.

Sealey, BA; James, LS; Cohen, G; Ryan, MJ; Page, RA. in press. Rapid foraging risk-assessments in the Jamaican fruit eating bat, Artibeus jamaicensis. Animal Behaviour.

Wilhite, K.; Ryan, M.J. 2024. Condition dependence in the sexual communication system of the túngara frog. Behavioral Ecology and Sociobiology. 78:54


Ryan, M.J. 2023. Sexual selection and the animal’s mating mind. pp. 16-32. In: D.M. Buss, editor, The Oxford Handbook of Human Mating. Oxford University Press, Oxford.

Dixon, M.M.; Carter, G.; Ryan, M.J.; Page, R.A. 2023. Spatial learning overshadows learning odors and sounds in both predatory and frugivorous bats. Behavioral Ecology.


Ryan, M.J. 2022. Sexual selection: Aesthetic appreciation and mate choice. pp. 218-239.  In: M. Skov & M. Nadal editors, The Routledge International Handbook of Neuroaesthetics. Routledge.

Ryan, M.J. 2022. Evolution of behavior. pp. 397-426. In: J.J. Bolhuis,  L.A. Giraldeau, J.A. Hogan, editors, The Behavior of Animals, Mechanisms, Function, and Evolution. John Wiley & Sons, Hoboken, New Jersey. [to purchase]

Ryan, M.J. 2022. Secret worlds: the extraordinary senses of animals. [book review] The Quarterly Review of Biology 97:55.

Christiano, B.M.; Ryan, M.J. 2022.  Diet of Brazilian Free-Tailed Bats (Tadarida brasiliensis): A review. Southwestern Naturalist 67:158-162. DOI:10.1894/0038-4909-67.2.158

Coss, D.; Ryan, M.J.; Page, R.A.; Hunter, K.L.; Taylor, R.C. 2022. Can you hear/see me? Multisensory integration of multimodal noise in túngara frogs. Behavioral Ecology

Dixon, M.M.; Jones, P.L.; Ryan, M.J.; Carter, G.C.; Page, R.A. 2022. Long-term memory in frog-eating bats. Current Biology

James, L.S.; Baier, A.L.; Page, R.A.; Clements, P.; Hunter, K.L.; Taylor, R.C.; Ryan, M.J. 2022. Cross-modal facilitation of auditory discrimination in a frog.  Biology Letters

James, L.S.; Taylor, R.C.; Hunter, K.L.; Ryan, M.J. 2022. Evolutionary and allometric insights into anuran auditory sensitivity and morphology.  Brain, Behavior and Evolution  DOI: 10.1159/000521309

Larter, L.C.; Bernal, X.E.; Page, R.A.; Ryan, M.J. 2022. Local competitive environment and male condition influence within-bout calling patterns in túngara frogs. Bioacoustics 10.1080/09524622.2022.2070544

Mendelson, T.C.; Ryan, M.J. 2022. Sex and design in our evolutionary cousins: The perception of beauty in nature. Metode Science Studies Journal

Ponnath, A; Ryan, M.J.; Fang, Z.; Farris, H.E. 2022.  Tuned in to the information bearing element in communication sounds: single cell sensitivity in the túngara frog midbrain to frequency modulated signals. PLoS ONE pone.0268383

Rosenthal, G.G.; Ryan, M.J. 2022. The ascent of women: Mate choice research since Darwin. Science 375, eabi6308. 

Zhu, B-C.; Yang, Y.; Zhou, Y.; Deng, K.; Wang, T-L.; Wang, J-C.; Tang, Ye-Z.; Ryan, M.J.; Cui, J. 2022. Multisensory integration facilitates perceptual restoration of an interrupted call in a frog. Behavioral Ecology


Ryan, M.J. 2021. Darwin, sexual selection, and the brain. Proceedings of the National Academy of Sciences. DOI 10.1073/pnas.2008194118

Ryan, M.J. 2021. Resolving the problem of sexual beauty. pp. 162-182. In: J.M. DeSilva, editor, A Most Interesting Problem. What Darwin’s Descent of Man Got Right and Wrong About Human Evolution. Princeton University Press, Princeton.

Connelly, A.D.; Ryan, M.J. 2021. Phenotypic variation in an asexual-sexual fish system: visual lateralization. Frontiers in Ecology and Evolution

Hemingway, C.T.; Aversa, J.; Ryan, M.J.; Page, R.A. 2021. Context-dependent preferences in wild fruit bats, Animal Behaviour 179:65-72

James, L.S.; Halfwerk, W.; Hunter, K.L.; Page, R.A.; Taylor, R.C.; Wilson, P.S.; Ryan, M.J. 2021.  Covariation among multimodal components in the túngara frog’s courtship display. Journal of Experimental Biology 224, jeb241661

Leslie, C.E.; Walkowski, W.; Rosencrans, R.F.; Gordon, W.C.; Bazan, N.G.; Ryan, M.J.; Farris, H.E. 2021. Estrogenic modulation of retinal sensitivity in reproductive female túngara frogs.  Integrative and Comparative Biology

Schlaepfer, M.A.; Caldwell, D.; Rorabaugh, J.C.; Ryan, M.J.; Stump, S.; Sredl, J. 2021.  The use of evoked vocal responses to detect cryptic, low-density frogs in the field. Journal of Herpetology 55: 174-180.

Taylor, R.C.; Wilhite, K.O.; Ludovici, R.J.; Mitchell, K.M.; Halfwerk, W.; Page, R.A.; Ryan, M.J.; Hunter, K.L. 2021. Complex sensory environments alter mate choice outcomes. Journal of Experimental Biology doi:10.1242/jeb.233288

Yin, W.;  Xue, Q.; Tian, B.; Yang, S.; Li, Z.; Chen, Z.; Ryan, M.J.; Hoffmann, A.A. 2021. Flexible habitat choice by aphids exposed to multiple cues reflecting present and future benefits. Behavioral Ecology 32:286-296.

Zhu, B.; Zhou, Y.; Yang, Y.; Deng, K.; Wang, T.; Wang, J.; Tang, Y.; Ryan, M.J.; Cui, J. 2021. Multisensory modalities increase working memory for mating signals in a treefrog. Journal of Animal Ecology 90:1455-1465


Brenowitz, E.A.; Ryan, M.J.; Zakon, H.H. 2020. Dr. Walter Wilczynski, 1952 – 2020. Brain Behavior and Evolution DOI: 10.1159/000510074 

Goutte, S.; Muñoz, M.I.; Ryan, M.J.; Halfwerk, W. 2020. Floating frogs sound larger: environmental constraints on signal production drives call frequency changes. The Science of Nature

Hemingway, C.T.; Ryan, M.J.; Page, R.A. 2020. State-dependent learning influences foraging behaviour in an acoustic predator. Animal Behaviour 163:33-38

James, L.S.; Ryan, M.J. 2020. Perspectives regarding future experiments on categorical perception: a comment on Green et al. Behavioral Ecology.

Jones, P.L.; Divoll, T.; Dixon, M.M.; Aparicio, D.; Cohen, G.; Mueller, U.; Ryan, M.J.; Page, R.A. 2020. Sensory ecology of the frog-eating bat, Trachops cirrhosus, from DNA metabarcoding and behavior. Behavioral Ecology

Kime, N.M.; Goutte, S.; Ryan, M.J. 2020. Arginine vasotocin affects vocal behavior but not selective responses to conspecific calls in male túngara frogs. Hormones and Behavior

Leslie, C.E.; Rosencrans, R.F.; Walkowski, W.; Gordon, W.C.; Bazan, N.G.; Ryan, M.J.; Farris, H.  2020. Reproductive state modulates retinal sensitivity to light in female túngara frogs. Frontiers in Behavioral Neuroscience doi: 10.3389/fnbeh.2019.00293

Lynch, K.S.; Ryan, M.J. 2020. Understanding the role of incentive salience in sexual decision-making. Integrative and Comparative Biology  pp. 1-10, DOI:10.1093/icb/icaa094


Ryan, M.J.; Akre, K.A.; Baugh, A.T.; Bernal, X.E.; Lea, A.L.; Leslie, C.; Still, M.B.; Wylie, D.; Rand, A.S. 2019. Nineteen years of consistently positive and strong female mate preferences despite individual variation. The American Naturalist  194:125-134.

Ryan, M.J.; Page, R.A.; Hunter, K.L.; Taylor, R.C. 2019.  “Crazy love”—nonlinearity and irrationality in mate choice. Animal Behaviour 147:189-198 

Cronin, A.D.; Ryan, M.J.; Page, R.A.; Hunter, K.L.; Taylor, R.C.  2019. Environmental heterogeneity alters mate choice behavior for multimodal signals. Behavioral Ecology and Sociobiology 73, 43

Garcia, M.J.; Rodríguez-Brenes, S.; Kobisk, A.; Adler,  L.; Ryan, M.J.; Taylor, R.C.; Hunter, K.L. 2019. Epigenomic changes in the túngara frog (Physalaemus pustulosus): Possible effects of introduced fungal pathogen and urbanization. Evolutionary Ecology

Hemingway, C.T.; Lea A.M., Page, R.A.; Ryan, M.J. 2019Effects of information load on response times in frogs and bats: mate choice vs. prey choice. Behavioral Ecology and Sociobiology

Hemingway, C.T., Ryan, M.J., Page, R.A. 2019. Transitive foraging behavior in the frog–eating bats. Animal Behaviour  154:47-55.

McClelland, B.E.; Ryan, M.J.; Wilczynski, W. 2019. Does sexual dimorphism vary by population? Laryngeal and ear anatomy in cricket frogs (Acris crepitans). Current Zoology.

Still, M.B.; Lea, A.M.; Hofmann, H.A.; Ryan, M.J. 2019. Multimodal stimuli regulate reproductive behavior and physiology in male túngara frogs. Hormones and Behavior

Smit, J.A.H.; Loning, H.; Ryan, M.J.; Halfwerk, W. 2019. Environmental constraints on  size–dependent signalling affects mating and rival interactions. Behavioral Ecology.

Taylor, R.C.; Akre, K.A.; Wilczynski, W; Ryan, M.J. 2019. Behavioral and neural auditory thresholds in a frog. Current Zoology.


Ryan, M.J. 2018. A Taste for the Beautiful: The Evolution of Attraction. Princeton University Press, Princeton, New Jersey. [to purchase] [translations in Spanish, Japanese, and Korean are available]

Ryan, M.J. 2018.  Animal vocal communication: assessment and management roles. second edition. [book review]. Quarterly Review of Biology 93:39.

Baugh, A.T.; Gridi-Papp, M.; Ryan, M.J. 2018. A laryngeal fibrous mass is essential to the attractiveness of a multicomponent call in túngara frogs (Physalaemus pustulosus). Bioacoustics 27:231-243.

Halfwerk, W.; Blaas, M.; Kramer, L.; Hijner, N.; Trillo, P.A.; Bernal, X.E.; Page, R.A.; Goutte, S; Ryan, M.J.; Ellers, J. 2018. Adaptive changes in sexual signaling in response to urbanization. Nature Ecology and Evolution

Hemingway, C.T.; Ryan, M.J.; Page, R.A. 2018. Cognitive constraints on optimal foraging in frog-eating bats. Animal Behaviour 143:43-50.

Kime, N.M.; Ryan, M.J.; Wilson, P.S. 2018. Modelling the production of complex calls in the túngara frog (Physalaemus pustulosus). Bioacoustics

Salas, A.K.; Wilson, P.S.; Ryan, M.J. 2018.  Acoustic communication in the Bocon toadfish (Amphichthys cryptocentrus).   Environmental Biology of Fishes 101:1175-1193.


Ryan, M.J.; Jordan, L.J. 2017. Courtship and mate choice. In: Call, J., editor-in-chief, APA Handbook of Comparative Psychology: Vol. 1. Basic Concepts, Methods, Neural Substrate, and Behavior. Chapter 37.35

Baugh A.T.; Ryan, M.J. 2017. Vasotocin induces sexually dimorphic effects on acoustically-guided behavior in a tropical frog. Journal of Comparative Physiology A

Farris, H.E.; Ryan, M. J. 2017. Schema versus primitive perceptual grouping: The relative weighting of sequential versus spatial cues during an auditory grouping task in frogs. Journal of Comparative Physiology A 203:175-182.

Gomes, D.G.E.; Halfwerk, W.; Taylor, R.C.; Ryan, M.J.; Page, R.A. 2017. Multimodal weighting differences by bats and their prey: probing natural selection pressures on sexually selected traits. Animal Behavior 134:99-102

Halfwerk, W.; Smit, J.; Loning, H.; Lea, A.M.; Geipel, I.; Ellers, J.; Ryan, M.J. 2017. Environmental conditions limit attractiveness of a sexual signal. Nature Communications DOI: 10.1038/s41467-017-02067-1

Hemingway, C.T.; Ryan, M.J.; Page, R.A.  2017. Rationality in decision-making in the fringe-lipped bat, Trachops cirrhosus. Behavioral Ecology and Sociobiology. 71:94. doi:10.1007/s00265-017-2321-5 

Laverde-R, O.; Ryan, M.J.; Cadena, D.  2017. Evolution of bird communication signals: transference between signals mediated by sensory drive. bioRxiv, 142463.

Taylor, R.C.; Page, R.A.; Klein, B.A.; Ryan, M.J.; Hunter, K.L. 2017. Perceived synchrony of frog multimodal signal components is influenced by content and order. Integrative and Comparative Biology 57:902-909.

Venetar, K.; Ryan, M.J.; Wilczynski. 2017. Responses of male cricket frogs (Acris crepitans) to attenuated and degraded advertisement calls. Ethology


Baugh, A.T.; Ryan, M.J.; Bernal, X.; Rand, A.S.; Bee, M.A. 2016. Female túngara frogs do not experience the continuity illusion. Behavioral Neuroscience 130:62-74.

Cui,J.; Song, X.; Zhu,B.; Fang, G.; Tang, Y.;  Ryan, M.J. 2016. Receiver discriminability drives the evolution of complex sexual signals by sexual selection. Evolution 70:922–927.

Gomes, D.G.E; Page, R.A.; Geipel, I; Taylor, R.C.; Ryan, M.J.; W. Halfwerk. 2016. Bats perceptually weight prey cues across sensory systems when hunting in noise. Science 353:1227-1280. Suppl

Halfwerk, W.; Lea, A.; Guerra, M.; Page, R.A.; Ryan, M.J. 2016. Vocal responses to noise reveal the presence of the Lombard effect in a frog. Behavioral Ecology 27:669-676.

Halfwerk, W.; Ryan, M.J.; Wilson, P.S. 2016. Wind-and rain-induced vibrations impose different selection pressures on multimodal signaling. The American Naturalist 188:279-288.

Jansen, M.; Plath, M.; Brusquetti, F.; Ryan, M.J. 2016.  Asymmetric shift in advertisement calls of sympatric frogs. Amphibia Reptilia DOI:10.1163/15685381-00003038

Kosch, T.A.; Bataille, A.; Didinger, C.; Eimes, J.E.; Rodríguez-Brenes, S.; Ryan, M.J.; Waldman, B. 2016. Major histocompatibility selection dynamics in pathogen-infected túngara frog (Physalaemus pustulosus) populations.  Biology Letters 12: 20160345.

Rodríguez-Brenes, S.; Rodriguez, D.; Ibáñez, R.; Ryan, M.J. 2016. Spread of amphibian chytrid fungus across lowland populations of túngara frogs in Panamá. PLoS ONE 11(5): e0155745. doi:10.1371/journal.pone.0155745

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


Ryan, M.J.; Leslie, C.; Ryan, E.S. 2015. Physalaemus pustulosus (túngara frog). Sexual communication. Herpetological Review 46:415-416.

Jones, P.L.; Ryan, M.J.; Chittka, L. 2015. The influence of past experience with flower reward quality on social learning in bumblebees. Animal Behaviour

Jordan, L.J.; Ryan, M.J. 2015. The sensory ecology of adaptive landscapes. Biology Letters 11: 20141054.

Lea, A.L.; Ryan, M.J. 2015. Irrationality in  mate choice revealed by túngara frogs.  Science 349:964-966. Suppl materials

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 B. doi: 10.1098/rspb.2015.1403.

Smith, C.C.; Harris, R.M.; Lampert, K.P.; Schartl, M.; Hofmann, H.A.; Ryan, M.J. 2015. Copy number variation in the melanocortin 4 receptor gene and alternative reproductive tactics the swordtail Xiphophorus multilineatus. Environmental Biology of Fishes DOI 10.1007/s10641-014-0234-


Ryan, M.J. 2014. When seeing is deceiving: a comment on Kelley and Kelley. Behavioral Ecology 25:273-466-467.

Ryan, M. J.; Guerra, M.A. 2014. The mechanism of sound production in túngara frogs and its role in sexual selection and speciation. Current Opinion in Neurobiology 28:54-59.

Ryan, M.J.; Taylor, R.C. 2014. Measures of mate choice: comment on Dougherty and Shuker.  Behavioral Ecology.

Akre, K.L.; Bernal, X.; Rand, A.S.; Ryan, M.J. 2014. Harmonic calls and indifferent females: no preferencs for human consonance in an anuran. Proceedings of the Royal Society B 281:20140986 oi:10.1098/rspb.2014098.

Guerra, M.; Ryan, M.J.; Cannatella, D.C. 2014. Ontogeny of sexual dimorphism in the larynx of the túngara frog: Physalaemus pustulosus. Copeia 2014:123-129.

Halfwerk, W.; Dixon, M.M.; Ottens, K.J.; 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. The Journal of Experimental Biology 217:3038-3044  doi:10.1242/jeb.107482

Halfwerk,W.; Page, R.A.; Taylor, R.C.; Wilson, P.S.; Ryan, M.J. 2014. Crossmodal comparisons of signal components allow for relative-distance assessment. Current Biology

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 343:413-416. (Suppl. Material)

Hauser, M.D.; Yang, C.; Berwick, R.; Tattersall, I.; Ryan, M.; Watumull, J.; Chomsky, N.; Lewontin, R. 2014. The mystery of language evolution.  Frontiers in Psychology 5, Article 401, doi: 10.3389/fpsyg.2014.00401

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


Ryan, M.J. 2013.  The importance of integrative biology to sexual selection and communication.  pp. 233-255. In: U. Stegmann, editor, Animal Communication Theory: Information and Influence. Cambridge University Press, Cambridge UK.

Ryan, M.J.; Cummings, M.E. 2013. Perceptual biases and mate choice.  Annual Review of Ecology, Evolution and Systematics 44:437-459. (Supplemental Material)

Adler, K.; Narins, P.M.; Ryan, M.J. 2013. Robert Capranica (1931-2012) and the science of anuran communication.  Herpetological Review 44:554-556.

Jones, P.L.; Ryan, M.J.; Flores, V.; Page, R.A. 2014. When to approach novel prey cues? Social learning strategies in frog-eating bats. Proceedings of the Royal Society London B. 280,

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. 199:279-283.

Kime, N.M.; Ryan, M.J.; Wilson, P.S. 2013.  A bond graph approach to modeling the anuran vocal production system. Journal of the Acoustical Society of America 133:4133-4144

Page, R.A; Ryan, M.J.; Bernal, X.E. 2013. Be loved, be prey, be eaten. pp. 123-154.In: Animal Behavior, vol 3. Case Studies: Integration and Application of Animal Behavior (ed., K. Yasukawa), New York: Praeger. 

Rog, S.; Ryan, M.J.; Mueller, U.; Lampert, K.P. 2013.  Evidence for morphological and genetic diversification of túngara frog populations on islands.  Herpetological Conservation and Biology 8:228-239.

Taylor, R.C.; Ryan, M.J. 2013. Interactions of multisensory components perceptually rescue túngara frog mating signals. Science 341:273-274. (Supplemental Material)


Baugh, A.T,; Hoke, K.L.; Ryan, M.J. 2012. Development of communication behaviour: Receiver ontogeny in tungara frogs and a prospectus for a behavioural evolutionary development. The Scientific World Journal doi10.1100/2012/680132.

Dawson, B.; Ryan M.J.2012. Evoked vocal responses change with experience in male Physalaemus pustulosus. Copeia 2012:678-682.

Dawson, B.; Ryan, M.J. 2012.  Female preferences are not altered by early acoustic experience in the neotropical frog Physalaemus pustulosus. Journal of Herpetology 46:535-538. 

Duarte-Guterman, P.; Ryan, M.J.; Tudeau, V. 2012. Developmental expression of sex steroid and thyroid hormone-related genes and their regulation by triiodothyronine in the gonad-mesonephros of a Neotropical frog, Physalaemus pustulosus. General and Comparative Endocrinology 177:195-204.

Duarte-Guterman, P.; Ryan, M.J.; Hogaman, N.S.; Trudaeu, V.L. 2012. Developmental profiles and thyroid regulation of brain transcripts in frogs: A species comparison with emphasis on Physalaemus pustulosus. Brain, Behavior and Evolution 79:8812.

Willis, P.M.; Rosenthal, G.G.; Ryan, M.J. 2012. An indirect cue of predation risk counteracts female preference for conspecifics in a naturally hybridizing fish Xiphophorus birchmanni. PLoS ONE 7:e34802. doi10.100/2012/680132


Ryan, M.J. 2011.  The brain as a source of selection on the social niche: Examples from the psychophysics of mate choice in túngara frogs. Integrative and Comparative Biology pp. 1–15 doi:10.1093/icb/icr065

Ryan, M.J. 2011. Replication in field biology: The case of the frog-eating bat. Science 334:1229-1230.

Ryan, M.J. 2011. Sexual selection.  In: Grzimek's Animal Life Encyclopedia. Pp. 179-185. Van Nostrand Reinhold Co., New York.

Ryan, M.J. 2011. Sexual selection: A tutorial from the túngara frog.  In: Losos, J.B., ed.  In Light of Evolution, Essays from the Laboratory and Field. pp. 185-203. Roberts and Company, Greenwood Village CO.

Ryan, M.J.; Wilczynski, W. 2011.  An Introduction to Animal Behavior, An Integrative Approach. Cold Springs Harbor Laboratory Press, Cold Springs Harbor, New York. [to purchase]

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 333:752-753. [Supplemental Material]

Akre, K.L.; Ryan, M.J. 2011. Female túngara frogs elicit more complex mating signals from males.  Behavioral Ecology.

Baugh, A.T.; Ryan, M.J. 2011. The relative value of call embellishment in tungara frogs. Behaiovral Ecology and Sociobiology 65:359-367.

Bonochea, L.A.; Ryan, M.J. 2011. Localization error and search costs during mate choice in túngara frogs, Physalameus pustulosusEthology 116:56-62.

Bonochea, L.A.; Ryan, M.J. 2011. Predation risk increases permissiveness for heterospecific advertisement calls in túngara frogs, Physalaemus pustulosus. Animal Behaviour  82 347e352

Dapper,  A.L.; Baugh, A.T.; Ryan, M.J. 2011.  The sounds of silence as an alarm cue in túngara frogs. Biotropica 43: 380–385.

Farris,H.E.; Ryan, M.J. 2011. Relative comparisons enable auditory grouping in frogs. Nature Communications2:410 DOI:10.1038/ncoms1470|

O’Connell, L.A.; Matthews, B.J.; Ryan, M.J.; Hofmann, H.A. 2011.  Characterization of the dopamine system in brain of the túngara frog, Physalameus pustulosusBrain, Behavior and Evolution. DOI 10.1159/000321725

O'Connell, L.A.; Ding, J.H.; Ryan, M.J.; Hofmann, H.A. 2011. Neural distribution of the nuclear progesterone receptor in the túngara frog, Physalaemus pustulosus. Journal of Chemical Neuroanatomy. 41:137–147.

Rosenthal, G.G.; Ryan, M.J. 2011. Conflicting preferences within females: sexual selection versus species recognition. Biology Letters doi:10.1098/rsbl.2011.0027 

Smith, C.C.; Ryan, M.J. 2011. Strategy dependent plasticity in ejaculate traits.  Biology Letters

Taylor, R.C.; Klein, B.A.; Ryan, M.J. 2011. Inter-signal interaction and uncertain information in anuran multimodal signals. Current Zoology  57: 153−161.

Taylor, R.C.; Klein, B.A.; Stein, J.; Ryan, M.J. 2011. Multimodal signal variation in space and time: How important is matching a signal with its signaler?  Journal of Experimental Biology 214:815-820.

Willis, P.M.; Ryan, M.J.; Rosenthal G.G. 2011. Encounter rates with conspecific males influence  female mate choice in a naturally hybridizing fish.  Behavioral Ecology 22:1234-1240.


Ryan, M.J. 2010.  An improbable path. In: L. Drickamer, D. Dewsbury editors.  Leaders in Animal Behavior, The Second Generation. pp. 465-496. Cambridge University Press, Cambridge.

Ryan, M.J. 2010.  The family that works together stays together. [Dispatch]  Current Biology 20:R403-R404

Ryan, M.J. 2010. The túngara frog: A model for sexual selection and communication.  In: M.D.  Breed, J. Moore, eds., Encyclopedia of Animal Behavior. volume 3, pp. 453-461. Academic Press, Oxford.

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

Akre, K.L.; Ryan, M.J. 2010. Complexity increases working memory for mating signals. Current Biology 20:502-505.

Akre, K.L.; Ryan, M.J. 2010. Proximity-dependent response to variably complex mating signals in túngara frogs (Physalaemus pustulosus).  Ethology 116:1138-1145. 

Baugh, A. T. & Ryan, M. J. 2010. Ambient light alters temporal updating behaviour during mate choice in a Neotropical frog. Canadian Journal of Zoology 88:448–453.

Baugh, A.T. & Ryan, M.J. 2010. Mate choice in response to dynamic presentation of male advertisement signals in túngara frogs.  Animal Behaviour 79:145-152.

Baugh, A.T.; Ryan, M.J. 2010. The development of sexual behavior in túngara frogs. Journal of Comparative Psychology 124:60–88.

Baugh, A. T. & Ryan, M. J. 2010. Temporal updating during phonotaxis in male túngara frogs (Physalaemus pustulosus).  Amphibia Reptilia.31:449-454

Goutee, S.; Kime, N.M.; Argo, T.F. IV; Ryan, M.J. 2010. Calling strategies of male túngara frogs in response to dynamic playback. Behaviour 147:65-83.

Hoke, K.L.; Ryan, M.J.; Wilczynski, W. 2010. Sexually dimorphic sensory gating drives behavioral differences in túngara frogs. Journal of Experimental Biology 213:3463-3472

Kime, N.M.; Whitney, T.K.; Ryan, M.J.; Rand, A.S.; Marler, C.A. 2010. Treatment with arginine vasotocin alters mating calls and decreases attractiveness in male túngara frogs. General and Comparative Endocrinology 165:221-228.

Lampert , K.P.; Schmidt, C.; Fischer, P.; Volff, J-N.; Hoffmann, C.; Muck, J.; Lohse, M.J.; Ryan, M.J.; Manfred Schartl, M. 2010. Determination of onset of sexual maturation and mating behavior by melanocortin receptor 4 polymorphisms. Current Biology 20:1729-1734.

Owren, M.J.; Rendall, D.; Ryan, M.J. 2010. Redefining animal signaling: Influence versus     information in communication. Biology and Philosophy 25:755–780

Pfennig, K.S; Ryan, M.J. 2010. Evolutionary diversification of mating behaviour: using artificial neural networks to study reproductive character displacement and speciation. pp. 187-214.  In: C. Tosh and G. Ruxton, editors, Modeling Perception with Artificial Neural Networks, Cambridge University Press, Cambridge, UK.

Pröhl, H.; Ron, S.R.; Ryan, M.J. 2010. Ecological and genetic divergence between two lineages     of Middle American túngara frogs Physalaemus (=Engystomops) pustulosus. BMC Evolutionary Biology 10:146 (18 pp)

Shah, A.A.; Ryan, M.J.; Bevilaqua; Schalaepfer, M.A. 2010.  Prior experience alters the behavioral response of prey to a nonnative predator.  Journal of Herpetology 44:185-192.

Smith, C.C.; Ryan, M.J. 2010. Evolution of sperm quality but not quantity in the internally fertilized fish Xiphophorus nigrensisJournal of Evolutionary Biology 23:1759-1771.

Wilczynski, W.; Ryan, M.J. 2010.  The behavioral neuroscience of anuran social signal processing.  Current Opinion in Neurobiology 20:754-763.


Ryan, M.J. 2009. Communication in frogs and toads. In: L.R. Squire editor. Encyclopedia of Neuroscience, volume 2, pp. 1159-1166. Academic Press: Oxford.

Ryan, M.J.; Akre, K.L.; Kirkpatrick, M. 2009. Cognitive mate choice.  In: R. Dukas, J. Ratcliffe editors. Cognitive Ecology II. pp. 137-155. University of Chicago Press, Chicago. 

Baugh, A. T. & Ryan, M. J. 2009. Female túngara frogs vary in commitment to mate choice decisions. Behavioral Ecology. 20:1153-1159.

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. Behavioral Ecology and Sociobiology 63 1269-1279.

Bernal, X.; Page, R.A.; Argo, T.F. IV; Ryan, M.J.; 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.

Bernal, X.E.; Rand, A.S.; Ryan. M.J. 2009.  Task differences underlie sexual dimorphism in mating behaviour in túngara frogs, Physalaemus pustulosus. Proceedings of the Royal Society, London Series B. 276, 1323–1329.

Dawson, B.; Ryan, M.J. 2009. Early experience leads to changes in the advertisement calls of male Physalaemus pustulosus. Copeia 2009:221-226.

Funk, C.; Ryan, M.J.; Cannatella, D.C. 2009. Genetic divergence is more tightly related to call variation than landscape features in the Amazonian frogs Physalaemus petersi and P. freibergi. Journal of Evolutionary Biology 22:1839-1853.

Rendall, D.; Owren, M.J.; Ryan, M.J. 2009.  What do animal signals mean? Animal Behaviour 78:233-240.

Romero-Carvajal, A.; Saenz-Ponce, N.; Venegas-Ferrin, M.; Almedia-Reinoso, D.; Lee, C.; Bond, J.; Ryan, M.J.; Wallingford, J.B.; Delpino, E.M. 2009. Embryogenesis and laboratory maintenance of the foam-nesting túngara frogs, genus Engystomops (=Physalaemus). Developmental Dynamics 238:1444-1454.

Sosa, J.A.; Ryan, M.J.; Schlaepfer, M.A. 2009. Induced morphological variation in Lowland Leopard Frog larvae (Rana yavapainsis) does not confer a survival advantage against Green Sunfih (Lepomis cyanellus). Journal of Herpetology 43:460-468.

Summers, K.; Roney, K.; da Silva, J.; Capraro, G.; Cuthbertson, B.J.; Kazianis, S.; Rosenthal, G.G.; Ryan, M.J.; McConell, T.J. 2009.  Divergent patterns of selection on the DAB and DXB MHC class II loci in Xiphophorus fishes.  Genetica 135:379-390.


Baugh, A.T.; Akre, K.L.; Ryan, M.J.  2008.  Categorical perception of a natural, multivariate signal: mating call recognition in túngara frogs. Proceedings of the National Academy of Sciences, USA 105:8985-8988.

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.  Journal of Experimental Biology 211:1203-1210.

Engeszer, R.E.; Wang, G.; Alberici da  Barbiano, L.; Ryan, M.J.; Parichy, D.M. 2008.  Sex-specific perceptual spaces for a vertebrate basal social aggregative behavior. Proceedings of the National Academy of Sciences, USA 105:929-933.

Funk, C.W.; Angulo, A.; Caldwell, J.P.; Ryan, M.J.; Cannatella, D.C. 2008. Comparison of morphology and calls of two cryptic species of Physaleamus (Anura: Leptodactylidae). Herpetologica. 64:290-304.

Heubel, K.U.; Hornhardt, K.; Ollmann, T.; Parzefall, J.; Ryan, M.J.; Schlupp, I. 2008. Geographic variation in female mate-copying in the species complex of a unisexual fish, Poecilia formosa. Behaviour 148:1041-1064.

Hoke, K.L.; Ryan, M.J.; Wilczynski, W. 2008. Candidate neural locus for sex differences in reproductive decisions.  Biology Letters 4:518-521.

McLennan, D.A; Ryan, M.J. 2008. Female swordtails prefer the scent of heterospecific males (Xiphophorus continens: Poeciliidae). Animal Behaviour 75:1731-1737. 

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

Taylor, R.C.; Klein, B.A.; Stein, J.; Ryan, M.J. 2008.  Faux frogs: Multicomponent signalling and the value of robotics in animal behaviour. Animal Behaviour 76:1089-1097.


 Ryan, M.J. 2007. Sensory ecology: See me, hear me. [Dispatch]  Current Biology 17:1019-1021.

Ryan, M.J.; Akre, K.L.; Kirkpatrick. M. 2007. Mate choice [Primer]. Current Biology 17:313-316

Ryan, M.J.; Bernal, X.E.; Rand, A.S. 2007. Patterns of mating call preferences in túngara frogs, Physalaemus pustulosusJournal of Evolutionary Biology 20:2235-2247.

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

Bernal, X.; A.S. Rand; M.J. Ryan. 2007. Sex differences in response to non-conspecific advertisement calls: Receiver permissiveness in male and female túngara frogs.  Animal Behaviour 73:955-964.

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

Boul, K.E.; W.C. Funk; C.R. Darst; D.C. Cannatella; M.J. Ryan. 2007.  Sexual selection drives speciation in an Amazonian frog.  Proceedings of the Royal Society, London series B 274:399-406.

Engeszer, R.E.; da Barbiano, L.A.; Ryan, M.J.; Parichy, D.M. 2007. Timing and plasticity of shoaling behavior in the zebrafish, Danio rerio. Animal Behaviour 74: 1269-1275.

Hoke, K.L.; Ryan, M.J.; Wilczynski, W. 2007.  Integration of sensory and motor processing underlying social behaviour in túngara frogs.  Proceedings of the Royal Society, London B 274:641-649

Hoke, K. L.; M. J. Ryan; W. Wilczynski. 2007. Functional coupling between substantia nigra and basal ganglia homologs in amphibians.  Behavioral Neuroscience 121:393–1399

Lampert, K.P.; Bernal, X.E.; Rand, A.S.; Mueller, U.G.; Ryan, M.J. 2007. Physalaemus pustulosus: History influences genetic diversity and morphology.  Herpetologica 63:311-319.

Pfennig, K.S.; Ryan, M.J. 2007.  Character displacement and the evolution of mate choice: an artificial neural network approach.  Philosophical Transactions of the Royal Society 362:411-419.

Phelps, S.; Rand, A.S.; Ryan, M.J. 2007.  The mixed-species chorus as public information: Túngara frogs eavesdrop on a heterospecific.  Behavioral Ecology 18:108-114.

Schlaepfer, M.A.; Sredl, M.J.; Rosen, P.C.; Ryan, M.J. 2007. High prevalence of Batrachochytrium dendrobatidis in wild populations of lowland leopard frogs Rana yavapaiensis in Arizona. EcoHealth. 4:421-427.


Ryan, M.J. 2006.  Question and Answer with Michael J. Ryan.  Current Biology 16(24):1012-1013.

Ryan, M.J. 2006. Profile of: A. Stanley Rand (1932-2005).  Iguana 13:42-46.

Bernal, X.; Rand, A.S.; Ryan, M.J. 2006.  Acoustic preferences and localization performance of blood-sucking flies (Corethrella Coquillett).  Behavioral Ecology 709-715

Cummings, M.E.; García de León, F.J.; Mollaghan, D.M.; Ryan, M.J. 2006.  Is UV ornamentation an amplifier in swordtails?  Zebrafish 3:91-100.

Griddi Papp, M; Rand, A.S.; Ryan, M.J. 2006.  Complex call production in túngara frogs. Nature 441:38.

Hill, S.; Ryan, M.J. 2006.  The role of female quality in the mate choice copying behaviour of sailfin mollies.  Biology Letters 2:203-205.

Kirkpatrick, M.; Rand, A.S.; Ryan, M.J. 2006.  Mate choice rules in animals.  Animal Behaviour 72:1215-1225.

Lampert, K.P.; Bernal, X.; 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 pustulosusBehavioral Ecology and Sociobiology 59:796-804.

Lynch, K.S.; Crews, D.C.; Ryan, M.J.; Wilczyski, W. 2006.  Hormonal state influences aspects of female mate choice in the túngara frog (Physalaemus pustulosus).  Hormones and Behavior 49:450-457.

Page, R.A.; Ryan, M.J. 2006.  Social transmission of novel foraging behavior in bats: Frog calls and their referents.  Current Biology 16:1201-1205.

Phelps, S.M.; Rand, A.S.; Ryan, M.J. 2006.  A cognitive framework for mate choice and species recognition.  The American Naturalist 167:28-42.

Pauly, G.B.; Bernal, X.E.; Rand, A.S.; Ryan, M.J. in press.  The vocal sac increases call rate in the túngara frog, Physalaemus pustulosusComparative Physiology and Biochemistry 79:708-719

Pfennig, K.S.; Ryan, M.J. 2006.  Reproductive character displacement generates reproductive isolation among conspecific populations: an artificial neural network study.  Proceedings of the Royal Society, London B 273:1361-1368.

Pröhl, H.; Koshy, R.; Mueller, U.; Rand, A.S.; Ryan, M.J. 2006.  Geographic variation and behavioral traits in túngara frogs in a zone of secondary contact.  Evolution 60:1669-1679.


Ryan, M.J. 2005. Evolution of behavior. pp. 294-314. In: J.J. Bolhuis, L-A. Giraldeau, editors, The Behavior of Animals; Mechanisms, Function, and Evolution. Blackwell Pub. Co., Oxford.

Ryan, M.J. 2005. The evolution of behavior, and integrating it towards a complete and correct understanding of behavioral biology.  Journal of Animal Biology 55:419-439.

Ryan, M.J.; Cummings, M.E. 2005.  Animal signals and the overlooked costs of efficacy.  Evolution 59): 1160-1161.

Farris, H.E.; Rand, A.S.; Ryan, M.J. 2005.  The effect of time, space and spectrum on auditory grouping in túngara frogs.  Journal of Comparative Physiology 191:1173-1183.

Gabor, C.R.; Ryan, M.J.; Morizot, D.C. 2005.  Character displacement in sailfin mollies, Poecilia latipinna: allozymes and behavior.  Environmental Biology of Fishes 73:75-88.

Hoke, K.L.; Ryan, M.J.; Wilczynski, W. 2005.  Social cues shift functional connectivity in the hypothalamus.  Proceedings of the National Academy of Sciences, USA 102:10712-10717.

Lynch, K.S.; Rand, A.S.; Ryan, M.J.; Wilczynski, W. 2005.  Plasticity in female mate choice associated with changing reproductive states.  Animal Behaviour 69:689-699.

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

Rosenthal, G.G.; Ryan, M.J. 2005.  Assortative preferences for stripes in danios.  Animal Behaviour 70:1063-1066.

Weight, L.A.; Crawford, A.J.; Rand, A.S.; Ryan, M.J. 2005.  Biogeography of the túngara frog, Physalaemus pustulosus: a molecular perspective.  Molecular Ecology 14:3857-3876

Witte, K.; Farris, H.E.; Ryan, M.J.; Wilczynski, W. 2005. How cricket frog females deal with a  noisy world: habitat-related differences in auditory tuning.  Behavioral Ecology 16:571-579.


Ryan M.J.2004.  Fickle females? [News & Views]. Nature 428:708-709.

Boul, K.E.; Ryan, M.J. 2004.  Population variation of complex advertisement calls in Physalaemus petersi and comparative laryngeal morphology.  Copeia 3:624-631.

Engeszer, R.E.; Ryan, M.J.; Parichy, D.M. 2004. Learned social preference in zebrafish. Current Biology 14:881-884.

Hoke, K.L.; Burmeister, S.S.; Fernald, R.D.; Rand, A.S.; Ryan, M.J.; Wilczynski, W. 2004.  Functional mapping of the auditory midbrain during mate call reception.  The Journal of Neuroscience 24:11264-11272.

Kime, N.M.; Burmeister, S.S.; Ryan, M.J. 2004.  Female preference for socially variable call characters in the cricket frog, Acris crepitansAnimal Behaviour 68:1391-1399.

Roney, K.E.; Cuthbertson, B.J.; Godwin, U.B.; Kazianis, S.; Coletta, L.D.; Rosenthal, G.G.; Ryan, M.J.; Schmidt, M.; McConnell, T.J. 2004.  Alternative splicing of major histocompatibility complex class II DXB transcripts in Xiphophorus fishes.  Immunogenetics 56:462-466. 

Rosenthal, G.G.; Rand, A.S.; Ryan, M.J. 2004.  The vocal sac as a visual cue in anuran communication: an experimental analysis using video playback.  Animal Behaviour 68:55-58.


Ryan, M.J.; Kime, N.M. 2003. Selection on long distance acoustic signals. pp. 225-274. In: Simmons, A.; Fay, R.; Popper, A. editors, Springer Handbook of Auditory Research; Acoustic Communication. Springer Verlag, Berlin.

Ryan, M.J.; Rand, A.S. 2003.  Mate recognition in túngara frogs: a review of some studies of brain, behavior, and evolution.  Acta Zoologica Sinica 49:713-726.

Ryan, M.J.; Rand, A.S. 2003. Sexual selection and female preference space: How female túngara  frogs perceive and respond to complex population variation in acoustic mating signals. Evolution 57:2608-2618.

Ryan, M.J.; Rand, W.; Hurd, P.L.; Phelps, S.M.; Rand. A.S. 2003. Generalization in response to mate recognition signals. The American Naturalist 161:380-394.

Castellano, S.; Giacoma, C.; Ryan, M.J. 2003. Call degradation in diploid and tetraploid green toads. Biological Journal of the Linnean Society 78:11-26.

Cummings, M.E.; Rosenthal, G.G.; Ryan, M.J. 2003. A private ultraviolet channel in visual communication. Proceedings of the Royal Society, London B 270: 897-904.

Kingston, J.; Rosenthal, G.G.; Ryan, M.J. 2003. The role of sexual selection in maintaining a colour polymorphism in the pygmy swordtail, Xiphophorus pygmaeus. Animal Behaviour 65:735-743.

Lampert, K.P.; Rand, A.S.; Mueller, U.G.; Ryan, M.J. 2003. Fine scale genetic pattern and evidence for sex-biased dispersal in the túngara  frog, Physalaemus pustulosus. Molecular Ecology 12:3325-3334.

Rosenthal, G.G. et al. 2003. Dissolution of sexual signal complexes in a hybrid zone between the swordtails Xiphophorus birchmanni and X. malinche (Poeciliidae). Copeia 2003:299-307.


.Autumn, K.; Ryan, M.J.; Wake, D.B. 2002. Integrating historical and mechanistic biology enhances the study of adaptation. Quarterly Review of Biology 77:383-408.

Bosch, J.; Rand, A.S.; Ryan, M.J. 2002. Response to variation in chuck frequency by male and female túngara frogs. Herpetologica 58:95-103.

Burmeister, S.S.; Ophir, A.G.; Ryan, M.J.; Wilczynski, W. 2002. Information transfer during cricket frog contests. Animal Behaviour 64:715-725.

Farris, H.F.; Rand,. A.S.; Ryan. M.J. 2002. The effects of spatially separated call components on phonotaxis in túngara frogs: Evidence for auditory grouping. Brain, Behavior and Evolution 60:181-188

Pröhl, H.; Adams, R.; U. Mueller; Rand, A.S.; Ryan, M.J. 2002. Polymerase chain reaction primers for polymorphic microsatellite loci from the túngara frog Physalaemus pustulosus. Molecular Ecology Notes 2:341-343.

Rosenthal, G.G.; Wagner, W.E. Jr.; Ryan, M.J. 2002. Secondary reduction of preference for swords in the pygmy swordtail Xiphophorus nigrensis (Pisces: Poeciliidae). Animal Behaviour 63:37-45.

Tárano, Z.; Ryan, M.J. 2002. No preexisting biases for heterospecific calls traits in the frog Physalaemus enesefae. Animal Behaviour 64:599-607.

Witte, K.; Ryan, M.J. 2002. Mate-choice copying in the sailfin molly (Poecilia latipinna) in the wild. Animal Behaviour 63:943-949.


Ryan, M.J. (editor). 2001. Anuran Communication. Smithsonian Institution Press, Washington D.C. [to purchae]

Ryan, M.J. 2001. Food, song and speciation. [News and Views]. Nature 409:139-140.

Ryan, M.J. 2001. Seeing red in speciation. [News & Views]. Nature 411:900-901.

Ryan, M.J.; Phelps S.M.; Rand A.S. 2001. How evolutionary history shapes recognition mechanisms. Trends in Cognitive Science 5:143-148.

Ryan, M.J.; Rand. A.S. 2001. Feature weighting in signal recognition and discrimination by the túngara frog. Pp. 86-101. In: Ryan, M.J. editor, Anuran Communication. Smithsonian Institution Press, Washington D.C.

Ryan, M.J.; Rosenthal, G.G. 2001. Variation and selection in swordtails. pp. 133-148. In: Dugatkin, L.A. editor, Model Systems in Behavioral Ecology. Princeton University Press, Princeton, New Jersey.

Dries L.A.; Morris M.R.; Ryan, M.J. 2001. Why are some male pygmy swordtails large? Copeia 2001: 355-364.

Gabor, C.R.; Ryan, M.J. 2001. Geographic variation in reproductive character displacement in mate choice by male sailfin mollies. Proceedings of the Royal Society, London B 268:1063-1071.

Phelps, S.M.; Rand, A.S.; Ryan, M.J. 2001. Vestigal preference functions in neural networks and túngara frogs. Proceedings of the National Academy of Sciences USA 98:13161-13166.

Rosenthal , G.G.; Flores Martinez, T.Y.; García de León, F.J.; Ryan, M.J. 2001. Shared preferences by predators and females for males ornaments in swordtails. The American Naturalist 158:146-154.

Schlupp, I.; McNab, R.; Ryan, M.J. 2001. Sexual harassment as a cost for molly females: bigger males cost less. Behaviour:138:277-286.

Wilczynski, W.; Rand A.S.; Ryan, M.J 2001. Evolution of calls and auditory tuning in the Physalaemus pustulosus species group. Brain, Behavior and Evolution 58:137-151.

Witte, K.; Ryan, M.J.; Wilczynski, W. 2001. Changes in the frequency structure of a mating call decreases its attractiveness to females in the cricket frog Acris creptians blanchardi. Ethology 107:685-699.


Ryan, M.J.; Getz, W. 2000. Signal decoding and receiver evolution: An analysis using an artificial neural network. Brain, Behavior and Evolution 56:45-62.

Bosch, J.; Rand, A.S.; Ryan, M.J. 2000. Acoustic competition in Physalaemus pustulosus, a differential response to calls of relative frequency. Ethology 106:865-871.

Bosch, J.; Rand , A.S. Ryan, M.J. 2000. Signal variation and call preferences for whine frequency in the túngara frog, Physalaemus pustulosus. Behavioral Ecology and Sociobiology 49:62-66.

Kime N.M.; Turner W.R; Ryan M.J. 2000. The transmission of advertisement calls in Central American frogs. Behavioral Ecology 11:71-83.

Marsh, D.M.; Rand, A.S.; Ryan M.J. 2000. Effects of inter-pond distance on breeding ecology of túngara frogs. Oecologia 122:505-513.

Phelps, S.M.; Ryan, M.J. 2000. History influences signal recognition: Neural network models of túngara frogs. Proceedings of the Royal Society, London B 267:1633-1639.

Rosenthal, G.G.; Ryan, M.J. 2000. Visual and acoustic communication in nonhuman animals: a comparison. Journal of Bioscience 25:285-290.

Sun, L.; Wilczynski, W.; Rand, A.S; Ryan, M.J. 2000. Trade-off in short and long distance communication in túngara (Physalaemus pustulosus) and cricket (Acris crepitans) frogs. Behavioral Ecology 11:102-109.

Witte, K.; Chen, K-C.; Wilczynski, W.; Ryan, M.J. 2000. Influence of amplexus on phonotaxis in the cricket frog Acris crepitans blanchardi. Copeia 2000:257-261.


Ryan, M.J. 1999. Phylogeny and mate choice. Mus. Reg. Sci. nat. Torino 1999:387-392.

Ryan, M.J. 1999. Sexual selection and sensory exploitation [Response]. Science 283:1083a

Ryan, M.J. 1999. Electrifying diversity [News & Views]. Nature 400:211-212.

Ryan, M.J.; Rand. A.S. 1999. Phylogenetic inference and the evolution of communication in túngara frogs. pp. 535-557. In: Hauser, M.; Konishi, M. editors, The Design of Animal Communication. MIT Press, Cambridge, MA.

Ryan, M.J.; Rand. A.S. 1999. Phylogenetic influences on mating call preferences in female túngara frogs (Physalaemus pustulosus). Animal Behaviour 57:945-956.

Burmeister, S.; Wilczynski, W.; Ryan, M.J. 1999. Temporal call changes and social context affect graded signaling in cricket frogs. Animal Behaviour 57:611-618 .

McLennan, D.A.; Ryan, M.J. 1999. Interspecific recognition and discrimination based upon olfactory cues in swordtails. Evolution 53:880-888.

Schlupp, I.; Waschulewski, M.; Ryan, M.J. 1999. Female preferences for naturally-occurring novel male traits. Behaviour 136; 519-527.

Wilczynski, W.; Rand, A.S.; Ryan, M.J. 1999. Female preferences for temporal order of call components in the túngara frog: A Bayesian analysis. Animal Behaviour 58:841-851.

Wilczynski, W.; Ryan, M.J. 1999. Geographic variation in animal communication systems. In: Foster, S.A.; Endler, J., editors, Geographic Diversification of Behavior: An Evolutionary Perspective, pp . 234-261. Oxford University Press, Oxford.


Ryan, M.J. 1998. Receiver biases, sexual selection and the evolution of sex differences. Science. 281:1999-2003.

Ryan, M.J. 1998. A model fish (review of Sex, Color and Mate Choice in Guppies). Trends in Ecology and Evolution 13:294.

Ryan, M.J. 1998. Principle with a handicap (review of The Handicap Principle). Quarterly Review of Biology 73477-479.

Ryan, M.J.; Autumn, K,; Wake, D.B. 1998. Integrative biology and sexual selection. Integrative Biology 1:68-72.

Ryan, M.J.; Kime, N.M.; Rosenthal, G.G. 1998. Patterns of evolution in human speech processing and animal communication. Behavioral and Brain Science 21:282-283.

Ryan, M.J.; Rand. A.S. 1998. Evoked vocal response in male túngara frogs: Preexisting biases in male responses? Animal Behaviour 56:1509-1516.

Cannatella, D.C.; Hillis, D.M.; Chippinendale, P.; Weigt, L.; Rand, A.S.; Ryan M.J. 1998. Phylogeny of frogs of the Physalaemus pustulosus species group, with an examination of data incongruence. Systematic Biology 47:311-335.

Kime, N.M., Rand, A.S., Kapfer, M., Ryan, M.J. 1998. Consistency of female choice in the túngara frog: A permissive preference for complex characters. Animal Behaviour 55:641-649.

McClelland, B.E.; Wilczynski, W.; Ryan, M.J. 1998. Intraspecific variation in laryngeal and ear morphology in male cricket frogs (Acris crepitans). Biological Journal of the Linnean Society 63:51-67.

Phelps, S.M.; Ryan, M.J. 1998. Neural networks predict response biases in female túngara frogs. Proceeding of the Royal Society, London B 265:279-285.

Witte, K.; Ryan, M.J. 1998. Male body size influences mate-choice copying in the sailfin molly Poecilia latipinna. Behavioral Ecology 9:534-539.


Ryan, M.J. 1997. Sexual selection and mate choice. pp. 179-202. In: J.R. Krebs, N.B. Davies, editors, Behavioural Ecology, An Evolutionary Approach. fourth edition. Blackwell, Oxford.

Ryan, M.J. 1997. Animal communication and evolution (review of Evolution and Communication). Evolution 51:1333-1337.

Marler, C.A.; Foran, C. Ryan, M.J. 1997. The influence of experience on mating preferences of the gynogenetic Amazon molly. Animal Behaviour 53:1035-1041.

Marler, C.A.; Ryan, M.J. 1997. Origin and maintenance of a female mating preference. Evolution 51:1244-1248.

McLennan, D.A.; Ryan, M.J. 1997. Responses to conspecific and heterospecific olfactory cues in the swordtail Xiphophorus cortezi. Animal Behaviour 54:1077-1088.

Rand, A.S.; Bridarolli, M.E.; Dries, L.; Ryan, M.J. 1997. Light levels influence female choice in túngara frogs: predation risk assessment? Copeia 1997:447-450.

Schlupp, I.; Ryan, M.J. 1997. Male mollies (Poecilia latipinna) copy the mate choice of other males. Behavioral Ecology 8:104-107.


Ryan, M.J. 1996. Phylogenetics and behavior: some cautions and expectations. pp. 1-21. In: Martins, E. editor, Phylogenies and the Comparative Method in Animal Behavior. Oxford University Press, Oxford.

Ryan, M.J. 1996. Speciation and the recognition concept, theory and application (review of). Quarterly Review of Biology 71:274-275.

Ryan, M.J. 1996. A natural history of amphibians (review of). Animal Behaviour 52:1058.

Ryan, M.J.; Dries, L.A.; Batra, P.; Hillis, D.M. 1996. Male mate preference in a gynogenetic species complex of Amazon mollies. Animal Behaviour 52:1225-1236.

Ryan, M.J.; Rand, A.S.; Weigt, L.A. 1996. Allozyme and advertisement call variation in the túngara frog, Physalaemus pustulosus. Evolution 50:2435-2453.

Marler, C.A.; Ryan, M.J. 1996. Energetic constraints and steroid hormone correlates of male calling behaviour in the túngara frog. Journal of Zoology 240:397-409.

McClelland, B.E.; Wilczynzksi, W.; Ryan, M.J. 1996. Correlations between call characteristics and morphology in male cricket frogs (Acris crepitans). Journal of Experimental Biology 199:1907-1919.

Morris, M.R.; Wagner, W.E. Jr.; Ryan, M.J. 1996. A negative correlation between trait and and mate preference in Xiphophorus pygmaeus. Animal Behaviour 52:1193-1203.

Morris, M.R.; Ryan, M.J. 1996. Sexual difference in signal-receiver coevolution. Animal Behaviour 52:1017-1024.

Schlupp, I.; Ryan, M.J. 1996. Mixed-species shoals and the maintenance of a sexual-asexual mating system in mollies. Animal Behaviour 52:885-890.


Ryan, M.J. 1995. Offsetting advantages (review of Sexual Selection). Science 267:712-713.

Ryan, M.J.; Rand, A.S. 1995. Female responses to ancestral advertisement calls in the túngara frog. Science 269, 390-392.

Morris, M.R.; Gass, L.; Ryan, M.J. 1995. Assessment and individual recognition of opponents in the pygmy swordtails Xiphophorus nigrensis and X. multilineatus. Behavioral Ecology 6:274-279.

Ryan. M.J.; Warkentin, K.M.; McClelland, B.E.; Wilczynski, W. 1995. Fluctuating asymmetries and advertisement call variation in the cricket frog, Acris crepitans. Behavioral Ecology 6:124-131.

Baer, C.F.; Dantzker, M.; Ryan, M.J. 1995. A test for preferece of association in a  color polymorphic poeciliid fish: laboratory study. Environmental Biology of Fishes 43:207-212.

Cocroft, R.C.; Ryan, M.J. 1995. Patterns of mating call evolution in toads and chorus frogs. Animal Behaviour 49:283-303.

Morris, M.R.; Gass, L.; Ryan, M.J. 1995. Assessment and individual recognition of opponents in the pygmy swordtails Xiphophorus nigrensis and X. multilineatus. Behavioral Ecology 6:274-279.

Morris, M.R.; Mussel, M.; Ryan, M.J. 1995. Vertical bars on male Xiphophorus multilineatus: a signal that deters rival males and attracts females. Behavioral Ecology 6:274-279.

Morris, M.R.; Ryan, M.J. 1995. Large body size in the pygmy swordtail Xiphophorus pygmaeus. Biological Journal of the Linnean Society 54:383-395.

Sullivan, B.K.; Ryan, M.J.; Verrill, P. 1995. Sexual selection and mating systems in amphibians. pp. 469-517. In: Sullivan, B.K.; Heatwole, H., editors, Social Behavior of Amphibians, Surrey Beatty & Sons, New York.

Wilczynski, W.; Rand, A.S.; Ryan, M.J. 1995. The processing of spectral cues by the call analysis system of the túngara frog, Physalaemus pustulosus. Animal Behaviour 49:911-929.


Ryan, M.J. 1994. Mechanistic studies in sexual selection. pp. 190-215. In: Real, L., editor, Behavioral Mechanisms in Evolutionary Ecology. University of Chicago Press, Chicago.

Foran, M.C.; Ryan, M.J. 1994. Female-female competition in a unisexual/bisexual complex of mollies. Copeia 1994:504-508.

Schlupp, I.; Marler, C.A.; Ryan, M.J. 1994. Benefit to male sail fin mollies of mating with heterospecific females. Science 263:373-374.


Ryan, M.J. 1993. Sexual selection on P-alleles and the evolution of mating asymmetries in swordtails (Xiphophorus nigrensis and X. pygmaeus). pp. 269-277. In: J.H. Schröder, J. Bauer, M. Schartl, editors, Trends in Ichthyology, An International Perspective. Blackwell Scientific Publications, Oxford.

Ryan, M.J.; Rand, A.S. 1993. Phylogenetic patterns of behavioral mate recognition systems in the Physalaemus pustulosus species group (Anura: Leptodactylidae): the role of ancestral and derived characters and sensory exploitation. pp. 251-267. In: D.R. Lees, D. Edwards, editors, Evolutionary Patterns and Processes. Linnean Society Symposium Series, No. 14, Academic Press, London.

Ryan, M.J.; Rand, A.S. 1993. Species recognition and sexual selection as a unitary problem in animal communication. Evolution 47:647-657.

Ryan, M.J.; Rand. A.S. 1993. Sexual selection and signal evolution: the ghost of biases past. Philosophical Transactions of the Royal Society B. 340:187-195.


Ryan, M.J. 1992. Sexy males and sterile females (review of The Peacock and the Ant). Cell 71:367-368.

Ryan, M.J. 1992. Costs of reproduction. In: Feder, M., editor, Physiology of the Amphibians. pp. 426-434. University of Chicago, Chicago.

Ryan, M.J.; Keddy-Hector, A. 1992. Directional patterns of female mate choice and the role of sensory biases. American Naturalist 139:S4-S35.

Ryan, M.J.; Pease, C.M.; Morris, M.R. 1992. A genetic polymorphism in the swordtail Xiphophorus nigrensis: testing the prediction of equal fitnesses. American Naturalist 139:21-31.

Ryan, M.J.; Perrill, S.A.; Wilczynski, W. 1992. Auditory tuning and call frequency predict population-based mating preferences in the cricket frog, Acris crepitans. American Naturalist 139:1370-1383.

Morris, M.J.; Batra, P.; Ryan, M.J. 1992. Male-male competition and access to females. Copeia 1992:980-986.

Morris, M.R.; Ryan, M.J. 1992. Breeding cycles in natural populations of Xiphophorus nigrensis, X. multilineatus and X. pygmaeus. Copeia 1992:1074-1077.

Keddy-Hector, A.; Wilczynski, W.; Ryan, M.J. 1992. Call patterns and basilar papilla tuning in cricket frogs. II. Intrapopulational variation and allometry. Brain, Behavior and Evolution 39:238-246.

Rand, A.S.; Ryan, M.J.; Wilczynski, W. 1992. Signal redundancy and receiver permissiveness in acoustic mate recognition by the túngara frog, Physalaemus pustulosus. American Zoologist 32:81-90.

Wilczynski, W.; Ryan, M.J. 1992. Introduction to the symposium: Mechanisms of mate choice. American Zoologist 32:15-17.

Wilczynski, W.; Keddy-Hector, A.; Ryan, M.J. 1992. Call patterns and basilar papilla tuning in cricket frogs. I. Differences among populations and between sexes. Brain, Behavior and Evolution 39:229-237.


Ryan, M.J. 1991. Sexual selection and communication in frogs: some recent advances. Trends in Ecology and Evolution 6:351-354.

Ryan, M.J.; Wilczynski, W. 1991. Evolution of intraspecific variation in the advertisement call of a cricket frog (Acris crepitans, Hylidae). Biological Journal of the Linnean Society 44: 249-271.

Kirkpatrick, M.; Ryan, M.J. 1991. The paradox of the lek and the evolution of mating preferences. Nature 350:33-38.


Ryan, M.J. 1990. Sensory systems, sexual selection, and sensory exploitation. Oxford Surveys in Evolutionary Biology 7:157-195.

Ryan, M.J. 1990. Signals, species, and sexual selection. American Scientist 78:46-52.

Ryan, M.J.; Cocroft, R.B.; Wilczynski, W. 1990. The role of environmental selection in intraspecific divergence of mate recognition signals in the cricket frog, Acris crepitans. Evolution 44:1869-1872.

Ryan, M.J.; Drewes, R.C. 1990. Vocal morphology of the Physalaemus pustulosus species group (Family Leptodactylidae): morphological response to sexual selection for complex calls. Biological Journal of the Linnean Society 40:37-52.

Ryan, M.J.; Fox, J.H.; Wilczynski, W.; Rand, A.S. 1990. Sexual selection for sensory exploitation in the frog Physalaemus pustulosus. Nature 343:66-67.

Ryan, M.J.; Hews, D.K.; Wagner, W.E. Jr. 1990. Sexual selection on alleles that determine body size in the swordtail Xiphophorus nigrensis. Behavioral Ecology and Sociobiology 26:231-237.

Ryan, M.J.; Rand, A.S. 1990. The sensory basis of sexual selection for complex calls in the túngara frog, Physalaemus pustulosus (sexual selection for sensory exploitation). Evolution 44:305-314.

Crapon de Caprona, M.; Ryan, M.J. 1990. Conspecific mate recognition in swordtails, Xiphophorus nigrensis and X. pygmaeus (Poeciliidae): olfactory and visual cues. Animal Behaviour 39:290-296.

Morris, M.R.; Ryan, M.J. 1990. Age at sexual maturity of Xiphophorus nigrensis in nature. Copeia 1990:745-751.


Ryan, M.J.; Causey, B.J. 1989. Alternative mating behavior in the swordtails Xiphophorus nigrensis and Xiphophorus pygmaeus (Pisces: Poeciliidae). Behavioral Ecology and Sociobiology 24:341-348.

Ryan, M.J.; Sullivan, B.K. 1989. Transmission effects on patterns of amplitude modulation in the advertisement calls of two toads, Bufo woodhousii and Bufo valliceps. Ethology 80:182-189.

Bruns, V.; Burda, H.; Ryan, M.J. 1989. Ear morphology of the frog-eating bat (Trachops cirrhosus, Family Phylostomidae): Apparent specializations for low-frequency hearing. Journal of Morphology 199:103-118.

Wilczynski, W.; Ryan, M.J.; Brenowitz, E.A. 1989. The display of the blue-black grassquit: the acoustic advantage of getting high. Ethology 80:218-222.


Ryan, M.J. 1988. Constraints and patterns in the evolution of anuran acoustic communication. In: B. Fritzsch, M. Ryan, W. Wilczynski, W. Walkowiak, T. Hetherington, editors, The Evolution of the Amphibian Auditory System. pp.637-677. John Wiley and Sons Inc., New York.

Ryan, M.J. 1988. Energy, calling, and selection. American Zoologist 28:885-898.

Ryan, M.J. 1988. Phenotype, genotype, swimming endurance and sexual selection in a swordtail (Xiphophorus nigrensis). Copeia 1988: 484-487.

Ryan, M.J.; Wilczynski, W. 1988. Coevolution of sender and receiver: effect on local mate preference in cricket frogs. Science 240:1786-1788.

Nyman, S.; Ryan, M.J.; Anderson, J.D. 1988. The distribution of the Ambystoma jeffersonianum complex in New Jersey, and a re-evaluation of some hypotheses on the origin and distribution of the complex. Journal of Herpetology 22:224-228.

Fritzsch, B.; Ryan, M.; Wilczynski, W.; Hetherington, T.; Walkowiak, W. (editors) 1988. The Evolution of the Amphibian Auditory System. John Wiley and Sons, New York [to purchase]

Wilczynski, W.; Ryan, M.J. 1988. The amphibian auditory system as a model for neurobiology, behavior and evolution. In: B. Fritzsch, M. Ryan, W. Wilczynski, W. Walkowiak, T. Hetherington, editors, The Evolution of the Amphibian Auditory System. pp. 3-12. John Wiley and Sons Inc., New York.


Ryan, M.J.; Tuttle, M.D. 1987. The role of prey generated sounds, vision, and echolocation in prey localization by the African bat Cardioderma cor (Megadermatidae). Journal of Comparative Physiology 161:59-66.

Ryan, M.J.; Wagner, W. 1987. Asymmetries in mating preferences between species: female swordtails prefer heterospecific mates. Science 236:595-597.


Ryan, M.J. 1986. Amphibians (review of Biology of Amphibians). Science 232:271.

Ryan, M.J. 1986. Environmental bioacoustics: evaluation of a commonly used experimental technique. Animal Behaviour 34:931-933.

Ryan, M.J. 1986. Factors influencing the evolution of acoustic communication: biological constraints. Brain, Behavior and Evolution 28:70-82.

Ryan, M.J. 1986. Neuroanatomy influences speciation rates among anurans. Proceedings of the National Academy of Sciences USA 83:1379-1382.

Ryan, M.J. 1986. The Panamanian love call. Natural History 95:36-43.

Ryan, M.J. 1986. Synchronized calling in a treefrog (Smilisca sila): short behavioral latencies and implications for neural pathways involved in call perception and detection. Brain, Behavior and Evolution 29:196-206.


Ryan, M.J. 1985. The Túngara Frog, A Study in Sexual Selection and Communication. University of Chicago Press, Chicago. 230 pp. [to purchase]

Ryan, M.J. 1985. Energetic efficiency of vocalization by the frog Physalaemus pustulosus. Journal of Experimental Biology 116:47-52.

Ryan, M.J.; Brenowitz, E.A. 1985. The role of body size, phylogeny, and ambient noise in the evolution of bird song. American Naturalist 126:87-100

Mendonca, M.; Licht, P.; Ryan, M.J.; Barnes, R. 1985. Changes in hormonal level in relation to breeding behavior in male bullfrogs (Rana catesbeiana) at the individual and population levels. General and Comparative Endocrinology 58:270-279.

Tuttle, M.D.; Ryan, M.J,; Bellwood, J. 1985. Acoustical resource partitioning by two phyllostomid bats. Animal Behaviour 33:1369-1371.


Ryan, M.J. 1983. Frequency modulated calls and species recognition in a Neotropical frog. Journal of Comparative Physiology 150:217-221.

Ryan, M.J. 1983. Sexual selection and communication in a Neotropical frog, Physalaemus pustulosus. Evolution 39:261-272.

Ryan, M.J.; Bartholomew, G.W.; Rand, A.S. 1983. Reproductive energetics of a Neotropical frog, Physalaemus pustulosus. Ecology 64:1456-1462.

Ryan, M.J.; Tuttle, M.D. 1983. The ability of the frog eating bat to distinguish among palatable and potentially poisonous prey using acoustic cues. Animal Behaviour 31:827-833

Ryan, M.J.; Tuttle, M.D.; Barclay, R.M.R. 1983. Behavioral responses of the frog-eating bat, Trachops cirrhosus, to sonic frequencies. Journal of Comparative Physiology 150:413-418.

Rand, A.S.; Ryan, M.J.; Troyer, K. 1983. A population explosion in a tropical frog: Hyla rufitela. Biotropica 15:72-73.

Waldman, B.; Ryan, M.J. 1983. Thermal advantages of communal egg deposition in Rana sylvatica. Journal of Herpetology 17:70-72.


Ryan, M.J. 1982. Variation in iguanine social organization: mating systems in chuckwallas (Sauromalus). In: G. Burghardt, A.S. Rand, editors, Iguanas of the World: Their Behavior, Ecology and Conservation. pp. 380-390. Noyes Press, Park Ridge, New Jersey.

Ryan, M.J.; Tuttle, M.D.; Rand, A.S. 1982. Sexual advertisement and bat predation in a Neotropical frog. The American Naturalist 119:136-139.

Bucher, T.L.; Ryan, M.J.; Bartholomew, G.W. 1982. Oxygen consumption during resting, calling and nest building in the frog Physalaemus pustulosus. Physiological Zoology 55:10-22.

Tuttle, M.D.; Ryan, M.J. 1982. The roles of synchronized calling, ambient noise, and ambient light in the anti-bat-predator behavior of a treefrog. Behavioral Ecology and Sociobiology 11:125-131.

Tuttle, M.D.; Taft, L.K.; Ryan, M.J. 1982. Acoustical location of calling frogs by philander opossums. Biotropica 13:233-234.

Tuttle, M.D.; Taft, L.K.; Ryan, M.J. 1982. Evasive behaviour of a frog in response to bat predation. Animal Behaviour 30:393-397.


Ryan, M.J.; Tuttle, M.D.; Taft, L.K. 1981. The costs and benefits of frog chorusing behavior. Behavioral Ecology and Sociobiology 8:273-278.

Barclay, R. M. R.; Fenton, B.; Tuttle, M. D.; Ryan, M. J. 1981. Echolocation calls produced by Trachops cirrhosus (Chiroptera: Phyllostomatidae) while hunting for frogs. Canadian Journal of Zoology 59:750-753

Rand, A.S.; Ryan, M.J. 1981. The adaptive significance of a complex vocal repertoire in a Neotropical frog. Zeitschrift fur Tierpsychologie 57:209-214.

Tuttle, M. D.; Ryan, M.J. 1981. Bat predation and the evolution of frog vocalizations in the Neotropics. Science 214:677-678.


Ryan, M.J. 1980. Female mate choice in a Neotropical frog. Science 209:523-525.

Ryan, M.J. 1980. The reproductive behavior of the bullfrog, Rana catesbeiana. Copeia 1980:108-114.


Ryan, M.J. 1978. A thermal property of the Rana catesbeiana (Amphibia, Anura, Ranidae) egg mass. Journal of Herpetology 12:295-297.

Ryan, M.J. 1978. Mirror image versus conspecific stimulation in adult male zebra finches. Wilson Bulletin 90:295-297.

Ryan, M.J. 1978. Parental care in salamanders. Bulletin of the New York Herpetological Society 13:23-27.

Anderson, J.D.; Hawthorne, K.; Galandak, J.; Ryan, M. 1978. A report on the endangered reptiles and amphibians of New Jersey. Bulletin of the New Jersey Academy of Science 23:26-33.

Prieto, A.A; Ryan, M.J. 1978. Some observations of the social behavior of the Arizona chuckwalla, Sauromalus obesus tumidus (Reptilia, Lacertilia, Iguanidae). Journal of Herpetology 12:327-336.


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