Publications

2026

Warkentin KM, Falk JJ, Casper AMA (2026) Sex across origins: Questioning animal-centric assumptions and developing integrative frameworks. Integr Comp Biol 66: icag150. https://doi.org.10.1093/icb/icag150

Eddy SL, Falk JJ, Massa MG, Warkentin KM, Casper AMA (2026) Teaching accurate sex-related biology: Approaches for more socially just instruction. Integr Comp Biol 66: icag136. https://doi.org/10.1093/icb/icag136

Casper AMA, Warkentin KM (2026) Synthesizing sex/es across scales and lineages: Centering plants, algae, fungi, protists, and modular metazoans. Integr Comp Biol 66:icag127. https://doi.org/10.1093/icb/icag127

 

Warkentin KM, Falk JJ, Casper AMA (2026) Evolution, development, and the incoherence of sex: A framework for multiple sex concepts. Integr Comp Biol 66: icag113. https://doi.org/10.1093/icb/icag113

2025

Guevara-Molina EC, Salazar-Nicholls MJ , Ribeiro Gomes F, Warkentin KM (2025) Ontogeny of the heat-induced hatching response in the red-eyed treefrog, Agalychnis callidryas. J Exp Zool A 345: 172181.https://doi.org/10.1002/jez.70049

Casper AMA, Fuselier LC, Jarosi S, Lewis AK, Ritz S, Warkentin KM, Jackson JK, Datta RR, Garriga-Lopez AM, Gordano S, Lane AK, Mark SL, Tanner KD, Zemenick AT, Eddy SL. (2025) Sustaining socially just and accurate life sciences teaching for sex, gender, and reproduction? Nat Sci Ed. 54:e70020. https://doi.org/10.1002/nse2.70020

 

Guevara-Molina EC, Ribeiro Gomes F, Warkentin KM (2025) Heat-induced hatching: clarifying effects of hydration and heating rate on behavioral thermal tolerance of red-eyed treefrog embryos. Integr Org Biol 7: obaf023. https://doi.org/10.1093/iob/obaf023

Jung J, Caldwell MS, McDaniel JG, Warkentin KM (2025) The role of vibration amplitude in the escape-hatching response of red-eyed treefrog embryos. Integr Org Biol 7: obaf012. https://doi.org/10.1093/iob/obaf012

Méndez-Narváez J, Warkentin KM (2025) Effects of larval foam-making and prolonged terrestriality on morphology, nitrogen excretion and development to metamorphosis in a leptodactylid frog. PeerJ. 13: e18990.https://doi.org/10.7717/peerj.18990

 

Salazar-Nicholls MJ, Macías Bazante H, Warkentin KM (2025) Functional morphology of hatching: Ontogeny and distribution of hatching gland cells in red-eyed treefrogs and a new marker for anuran hatching enzyme. J Morphol 286: e70029. https://doi.org/10.1002/jmor.70029

2024

Lisondro-Arosemena A, Salazar-Nicholls MJ, Warkentin KM (2024) Elevated ammonia cues hatching in red-eyed treefrogs: A mechanism for escape from drying eggs. J Exp Zool B 342: 406–411. https://doi.org/10.1002/jez.b.23253

– Media coverage including STRI (in English & Spanish), La Prenza,

Güell BA, McDaniel JG, Warkentin KM (2024) Egg-clutch biomechanics affect escape-hatching behavior and performance. Integr Org Biol 6: obae006. https://doi.org/10.1093/iob/obae006

2023

Güell BA, Warkentin KM (2023) To hatch and hatch not: does heterochrony in onset of vestibular mechanosensing explain species differences in escape-hatching success of Agalychnis embryos in snake attacks? Behav Ecol Sociobiol 77: 141. https://doi.org/10.1007/s00265-023-03417-4

 

Güell BA, Warkentin KM (2023) Phenology and environmental determinants of explosive breeding in gliding treefrogs: Diel timing of rainfall matters. Behav Ecol https://doi.org/10.1093/beheco/arad072

Méndez-Narváez J, Warkentin KM (2023) Early onset of urea synthesis and ammonia detoxification pathways in three terrestrially developing frogs. J Comp Physiol B 193: 523–543. https://doi.org/10.1007/s00360-023-01506-4

Gomez EK, Chaiyasarikul A, Güell BA, Warkentin KM (2023) Developmental changes in red-eyed treefrog embryo behavior increase escape-hatching success in wasp attacks. Behav Ecol Sociobiol 77: 52 https://doi.org/10.1007/s00265-023-03324-8

2022

Güell BA, Jung J, Almanzar A, Cuccaro-Díaz J, Warkentin KM (2022) Ontogeny of risk assessment and escape-hatching performance by red-eyed treefrog embryos in two threat contexts. J Exp Biol 225: jeb244533. DOI:10.1242/jeb.244533

Guevara-Molina et al 2022 IOB cover

Guevara-Molina EC, Ribeiro Gomes F, Warkentin KM (2022) Heat-induced hatching of red-eyed treefrog embryos: Hydration and clutch structure increase behavioral thermal tolerance. Integrative Organismal Biology obac041, DOI: 10.1093/iob/obac041

Majoris J, Francisco F, Burns C, Brandl S, Warkentin K, Buston P (2022) Paternal care regulates the timing, synchrony, and success of hatching in a coral reef fish. Proceedings of the Royal Society B 289: 20221466. DOI: 10.1098/rspb.2022.1466

Jung J, Guo M, Crovella ME, McDaniel JG, Warkentin KM (2022) Frog embryos use multiple levels of temporal pattern in risk assessment for vibration-cued hatching. Animal Cognition DOI: 10.1007/s10071-022-01634-4

Warkentin KM, Jung J, McDaniel JG (2022) Research approaches in mechanosensory-cued hatching and the iterative development of playback methods for red-eyed treefrog embryos. In Biotremology: Physiology, Ecology, and EvolutionAnimal Signals and Communication, vol. 8. Eds. Hill PSM, Mazzoni V, Stritih Peljhan N, Virant-Doberlet M, Wessel A.  Springer, Cham. p. 157–201. DOI: 10.1007/978-3-030-97419-0_7

Méndez-Narváez J, Warkentin KM (2022) Reproductive colonization of land by frogs: Embryos and larvae excrete urea to avoid ammonia toxicity. Ecology and Evolution DOI: 10.1002/ece3.8570

2021

Jung J, McDaniel JG, Warkentin KM (2021) Escape-hatching decisions show adaptive ontogenetic changes in how embryos manage ambiguity in predation risk cues. Behavioral Ecology and Sociobiology 75: 141. DOI: 10.1007/s00265-021-03070-9

Warkentin KM (2021) Queering herpetology: On human perspectives and the study of diverse animals. Chapter 4 in Herpetologia Brasileira Contemporânea. Org. Toledo LF. Sociedade Brazileira de Herpetologia, São Paulo. p 42–58 [Proceedings of the IX Brazilian Herpetology Congress: Inclusive Herpetology.] pdf file of chapter

González K, Warkentin KM, Güell BA (2021) Dehydration-induced mortality and premature hatching in gliding treefrogs with even small reductions in humidity. Ichthyology & Herpetology 109: 21–30. DOI: 10.1643/h2020085

Gomez EK, Warkentin KM, Güell BA (2021) Egg-kicking behaviour by male gliding treefrogs (Agalychnis spurrelli Boulenger, 1913) does not dislodge competitors’ eggs. Herpetology Notes 14: 157–161

2020

Jung J, Serrano-Rojas SJ, Warkentin KM (2020) Multimodal mechanosensing enables treefrog embryos to escape egg predators Journal of Experimental Biology 223: jeb236141. DOI: 10.1242/jeb.236141

Delia J, Bravo-Valencia L, Warkentin KM (2020) The evolution of extended parental care in glassfrogs: Do egg-clutch phenotypes mediate coevolution between the sexes? Ecological Monographs 90: e01411. DOI 10.1002/ecm.1411

 

 

2019

Jung J, Kim SJ, Pérez Arias SM, McDaniel JG, Warkentin KM (2019) How do red-eyed treefrog embryos sense motion in predator attacks? Assessing the role of vestibular mechanoreception. Journal of Experimental Biology 222: jeb206052. DOI 10.1242/jeb.206052

Warkentin KM, Jung J, Rueda Solano LA, McDaniel JG (2019) Ontogeny of escape-hatching decisions: vibrational cue use changes as predicted from costs of sampling and false alarms. Behav Ecol Sociobiol 73: 51. doi: 10.1007/s00265-019-2663-2

Cohen KL, Piacentino M, Warkentin KM (2019) Two types of hatching gland cells facilitate escape-hatching at different developmental stages in red-eyed treefrogs, Agalychnis callidryas (Anura: Phyllomedusidae). Biol J Linn Soc doi: 10.1093/biolinnean/bly214

Delia J, Rivera JM, Salazar Nicholls MJ, Warkentin KM (2019) Hatching plasticity and the adaptive benefits of extended embryonic development in glassfrogs. Evol Ecol 33: 37–53. doi: 10.1007/s10682-018-9963-2

2018

Güell BA, Warkentin KM (2018) When and where to hatch? Red-eyed treefrog embryos use light cues in two contexts. PeerJ doi: 10.7717/peerj.6018

Hite JL, Hughey MC, Warkentin KM, Vonesh JR (2018) Cross-ecosystem effects of terrestrial predators link treefrogs, zooplankton, and aquatic primary production. Ecosphere 9: e02377. doi: 10.1002/ecs2.2377

Cohen, KL, Piacentino ML, Warkentin KM. (2018) The hatching process and mechanisms of adaptive hatching acceleration in hourglass treefrogs. J Comp Biochem Physiol A. 217: 63–74 doi: 10.1016/j.cbpa.2017.10.020

2017

Salica MJ, Vonesh JR, Warkentin KM. (2017) Egg clutch dehydration induces early hatching in red-eyed treefrogs, Agalychnis callidryas. PeerJ. 5:e3549. doi: 10.7717/peerj.3549

Warkentin KM, Cuccaro Diaz J, Güell BA, Jung J, Kim SJ, Cohen KL. (2017) Developmental onset of escape-hatching responses in red-eyed treefrogs depends on cue type. Anim Behav. 129: 103–112. doi: 10.1016/j.anbehav.2017.05.008

Delia-JEB2017Cover-p150Delia J, Bravo-Valencia L, Warkentin KM. (2017) Patterns of parental care in Neotropical glassfrogs: fieldwork alters hypotheses of sex-role evolution. J Evol Biol. doi: 10.1111/jeb.13059

 

2016

Rueda Solano LA, Warkentin KM. (2016) Foraging behavior with possible use of substrate-borne vibrational cues for prey localization in Atelopus laetissimus (Ruiz-Carranza, Ardila-Robayo, and Hernández-Camacho, 1994). Herp Notes. 9: 191–195.

jexbio_219_12_cover.eps
Cohen KL, Seid MA, Warkentin KM. (2016) How embryos escape from danger: the mechanism of rapid, plastic hatching in red-eyed treefrogs. J Exp Biol 219: 1875-1883. doi: 10.1242/jeb.139519

Bouchard&al2016-FEcover-p150

Bouchard SS, O’Leary CJ, Wargelin LJ, Charbonnier JF, Warkentin KM (2016) Post-metamorphic carryover effects of larval digestive plasticity. Funct Ecol 30: 379–388 doi: 10.1111/1365-2435.12501

2015

Bouchard SS, O’Leary CJ, Wargelin LJ, Rodriguez WB, Jennings KX, Warkentin KM (2015) Alternative competition-induced digestive strategies yield equal growth but constrain compensatory growth in red-eyed treefrog larvae. J Exp Zool 323A: 778–788. doi: 10.1002/jez.1991

McCoy MW, Wheat SK, Warkentin KM, Vonesh JR (2015) Risk assessment based on indirect predation cues: Revisiting fine-grained variation. Ecol Evol doi: 10.1002/ece3.1552

Hughey MC, Rogge JR, Thomas K, McCoy MW, Warkentin KM (2015) Escape-hatching responses of individual treefrog embryos vary with threat level in wasp attacks: a mechanistic analysis. Behaviour doi:10.1163/1568539X-00003291

Touchon JC, McCoy MW, Landberg T, Vonesh JR, Warkentin KM (2015) Putting µ/g in a new light: plasticity in life-history switch points reflects fine-scale adaptive responses. Ecology 96: 2192–2202.

Tarvin&al2015-BiotropicaCover-150Tarvin RD, Silva Bermúdez C, Briggs VS, Warkentin KM (2015) Carry-over effects of size at metamorphosis in red-eyed treefrogs: higher survival but slower growth of larger metamorphs. Biotropica 47:218–226.


2014

Willink B, Palmer MS, Landberg T, Vonesh JR , Warkentin KM (2014) Environmental context shapes immediate and cumulative costs of risk-induced early hatching. Evol Ecol 28:103-116

2013

Touchon&al2013-OecologiaCoverTouchon JC, Jiménez RR, Abinette SH, Vonesh JR, Warkentin KM (2013) Behavioral plasticity mitigates risk across environments and predators during anuran metamorphosis. Oecologia 173:801-811


Gomez-Mestre I, Warkentin KM (2013) Risk-induced hatching timing shows low heritability and evolves independently of spontaneous hatching in red-eyed treefrogs. J Evol Biol 26:1079-1089

Touchon JC, McCoy MW, Vonesh JR, Warkentin KM (2013) Effects of plastic hatching timing carry over through metamorphosis in red-eyed treefrogs. Ecology 94:850-860

2012

Hughey MC, McCoy MW, Vonesh JR, Warkentin KM (2012) Spatial contagion shapes colonization dynamics of frogflies (Megaselia sp. nov.) on clutches of red-eyed treefrogs (Agalychnis callidryas). Biol Lett 8:887-889

Hughey MC, Nicolás A, Vonesh JR, Warkentin KM (2012) Wasp predation drives the assembly of fungal and fly communities on frog egg masses. Oecologia 168:1057-1068

McCoy MW, Touchon JC, Landberg T, Warkentin KM, Vonesh JR (2012) Prey responses to predator chemical cues: disentangling the importance of the number and biomass of prey consumed by predators. PLoS ONE 7:e47495

Wibowo E, Wassersug R, Warkentin K, Walker L, Robinson J, Brotto L, Johnson T (2012) Impact of androgen deprivation therapy on sexual function: A response. Asian J Andrology 14:793-794 doi:10.1038/aja.2012.60

2011

Gonzalez S, Touchon JC, Vonesh JR (2011) Interactions between competition and predation shape early growth and survival of two Neotropical hylid tadpoles. Biotropica 43:633-639

McCoy MW, Bolker BM, Warkentin KM, Vonesh JR (2011) Predicting predation through prey ontogeny using size-dependent functional response models. Am Nat 177:752-766

Touchon JC, Urbina J, Warkentin KM (2011) Habitat-specific constraints on induced hatching in a treefrog with reproductive mode plasticity. Behav Ecol 22:169-175

Touchon JC, Warkentin KM (2011) Thermally contingent plasticity: temperature alters expression of predator-induced color and morphology in a Neotropical treefrog tadpole. J Anim Ecol 80:79-88

ICB-51(1)CoverWarkentin KM (2011) Environmentally cued hatching across taxa: Embryos respond to risk and opportunity. Integr Comp Biol 51:14-25

Warkentin KM (2011) Plasticity of hatching in amphibians: Evolution, trade-offs, cues and mechanisms. Integr Comp Biol 51:111-127


2010

2010 Current Biology coverCaldwell MS, Johnston GR, McDaniel JG, Warkentin KM (2010) Vibrational signaling in the agonistic interactions of red-eyed treefrogs. Curr Biol 20:1012-1017

Caldwell MS, McDaniel JG, Warkentin KM (2010) Is it safe? Red-eyed treefrog embryos assessing predation risk use two features of rain vibrations to avoid false alarms. Anim Behav 79:255-260

Gomez-Mestre&al2010-JEvB-cover150Gomez-Mestre I, Saccocio VL, Iijima T, Collins EM, Rosenthal GG, Warkentin KM (2010) The shape of things to come: linking developmental plasticity to post-metamorphic morphology in anurans. J Evol Biol 23:1364-1373


Touchon JC, Warkentin KM (2010) Short- and long-term effects of the abiotic egg environment on viability, development and vulnerability to predators of a Neotropical anuran. Funct Ecol 24:566-575

2009

Caldwell MS, McDaniel JG, Warkentin KM (2009) Frequency information in the vibration-cued escape hatching of red-eyed treefrogs. J Exp Biol 212:566-575

Touchon JC, Warkentin KM (2009) Negative synergism of rainfall patterns and predators affects frog egg survival. J Anim Ecol 78:715-723

Warkentin KM, Caldwell MS (2009) Assessing risk: embryos, information, and escape hatching. In: Dukas R, Ratcliffe JM (eds) Cognitive ecology II. University of Chicago Press, Chicago, pp 177-200

2008

Gomez-Mestre I, Touchon JC, Saccocio VL, Warkentin KM (2008) Genetic variation in pathogen-induced early hatching of toad embryos. J Evol Biol 21:791-800

Gomez-Mestre I, Wiens JJ, Warkentin KM (2008) Evolution of adaptive plasticity: risk-sensitive hatching in neotropical leaf-breeding treefrogs. Ecol Monogr 78:205-224

JEB coverRogge JR, Warkentin KM (2008) External gills and adaptive embryo behavior facilitate synchronous development and hatching plasticity under respiratory constraint. J Exp Biol 211:3627-3635


PNAS Cover-Touchon&Warkentin150Touchon JC, Warkentin KM (2008) Reproductive mode plasticity: Aquatic and terrestrial oviposition in a treefrog. Proc Natl Acad Sci USA 105:7495-7499


Touchon JC, Warkentin KM (2008) Fish and dragonfly nymph predators induce opposite shifts in color and morphology of tadpoles. Oikos 117: 634-640.

2007

Gomez-Mestre I, Warkentin KM (2007) To hatch and hatch not: similar selective trade-offs but different responses to egg predators in two closely related, syntopic treefrogs. Oecologia 153:197-206

Warkentin KM (2007) Oxygen, gills, and embryo behavior: mechanisms of adaptive plasticity in hatching. Comp Biochem Phys A 148:720-731

Warkentin KM, Caldwell MS, Siok TD, D’Amato AT, McDaniel JG (2007) Flexible information sampling in vibrational assessment of predation risk by red-eyed treefrog embryos. J Exp Biol 210:614-619 [Inside JEB commentary]

2006

untitledGomez-Mestre I, Touchon JC, Warkentin KM (2006) Amphibian embryo and parental defenses and a larval predator reduce egg mortality from water mold. Ecology 87:2570-2581


Touchon JC, Gomez-Mestre I, Warkentin KM (2006) Hatching plasticity in two temperate anurans: responses to a pathogen and predation cues. Can J Zool 84:556-563

Vonesh & Warkentin Ecology coverVonesh JR, Warkentin KM (2006) Opposite shifts in size at metamorphosis in response to larval and metamorph predators. Ecology 87:556-562


Warkentin KM, Buckley CR, Metcalf KA (2006) Development of red-eyed treefrog eggs affects efficiency and choices of egg-foraging wasps. Anim Behav 71:417-425

Warkentin KM, Caldwell MS, McDaniel JG (2006) Temporal pattern cues in vibrational risk assessment by red-eyed treefrog embryos, Agalychnis callidryas. J Exp Biol 209:1376-1384 [Inside JEB commentary]

Warkentin KM, Gray RE, Wassersug RJ (2006) Restoration of satisfying sex for a castrated cancer patient with complete impotence: a case study. J Sex Marital Ther 32:389-399

2005

Warkentin KM (2005) How do embryos assess risk? Vibrational cues in predator-induced hatching of red-eyed treefrogs. Anim Behav 70:59-71

Warkentin KM, Gomez-Mestre I, McDaniel JG (2005) Development, surface exposure, and embryo behavior affect oxygen levels in eggs of the red-eyed treefrog, Agalychnis callidryas. Physiol Biochem Zool 78:956-966

Earlier

Warkentin KM (2002) Hatching timing, oxygen availability, and external gill regression in the tree frog, Agalychnis callidryas. Physiol Biochem Zool 75:155-164

Warkentin KM, Currie CC, Rehner SA (2001) Egg-killing fungus induces early hatching of red-eyed treefrog eggs. Ecology 82:2860-2869

Warkentin KM, Wassersug RJ (2001) Do prostaglandins regulate external gill regression in anurans? J Exp Zool 289:366-373

Warkentin KM (2000) Wasp predation and wasp-induced hatching of red-eyed treefrog eggs. Anim Behav 60:503-510

Warkentin KM (2000) Environmental and developmental effects on external gill loss in the red-eyed treefrog, Agalychnis callidryas. Physiol Biochem Zool 73:557-565

Warkentin KM (1999) The development of behavioral defenses: a mechanistic analysis of vulnerability in red-eyed tree frog hatchlings. Behav Ecol 10:251-262

Warkentin KM (1999) Effects of hatching age on development and hatchling morphology in the red-eyed treefrog, Agalychnis callidryas. Biol J Linn Soc 68:443-470

Warkentin KM (1995) Adaptive plasticity in hatching age: a response to predation risk trade-offs. Proc Natl Acad Sci USA 92:3507-3510

Ryan MJ, Warkentin KM, McClelland BE, Wilczynski W (1995) Fluctuating asymmetries and advertisement call variation in the cricket frog, Acris crepitans. Behav Ecol 6:124-131

Warkentin KM (1992) Microhabitat use and feeding rate variation in green frog tadpoles (Rana clamitans). Copeia 1992:731-740

Warkentin KM (1992) Effects of temperature and illumination on feeding rates of green frog tadpoles (Rana clamitans). Copeia 1992:725-730