Hanscom RJ, Hill JL, Patterson C, Marbach T, Sukumaran J, Higham TE, Clark RW. 2023. Cryptic behavior and activity cycles of a small mammal keystone species revealed through accelerometry: a case study of Merriam’s kangaroo rats (Dipodomys merriami). Movement Ecology. 11:72.

Lin JW, Liao CP, Chou CC, Clark RW, Tseng HY, Hsu JY, Huang WS. 2023. Loss of sea turtle eggs drives the collapse of an insular reptile community. Science Advances. 9:eadj7052.

Maag DW, Fancioli YZ, ‡Shaw N, ‡Sony AY, Castoe TA, Schuett GW, Clark RW. 2023. Hunting behavior and feeding ecology of Mojave rattlesnakes (Crotalus scutulatus), prairie rattlesnakes (Crotalus viridis), and their hybrids in southwestern New Mexico. Ecology and Evolution 13:e10683.

Hanscom RJ, Higham TE, Ryan DS, Clark RW. 2023. Ambush hunting in snakes: behavior, function, and diversity. In: Penning D, editor. Snakes: Morphology, Function, and Ecology. Nova Science Publishers, Inc. p. 279–311.

Clark RW, 2023. Grand challenges in foraging behavior and predator-prey interactions: next generation ethology in the Anthropocene. Frontiers in Ethology, 2:1304654.

Tetzlaff SJ, Vizentin‐Bugoni J, Sperry JH, Davis MA, Clark RW, Repp RA, Schuett GW, 2023. Fission–fusion dynamics in the social networks of a North American pitviper. Ecology and Evolution, 13:e10339.

Hanscom RJ, DeSantis DL, Hill JL, Marbach T, Sukumaran J, Tipton AF, Thompson ML, Higham TE, Clark RW. 2023. How to study a predator that only eats a few meals a year: high-frequency accelerometry to quantify feeding behaviours of rattlesnakes (Crotalus spp.). Animal Biotelemetry. 11(1):20.

Maag DW, Schuett GW, Clark RW. 2023. Reproduction and sexual receptivity in Crotalus scutulatus (Mohave Rattlesnake). Herp Rev. 54:133.

Hansom RJ, Clark RW. 2023. Diet and scavenging in Crotalus viridis. Herpetological Review. 54:311.


Gibert RG, Maag DW, Sanders LN, Clark RW. 2022. Investigating personality in vipers: individual rattlesnakes exhibit consistent behavioral responses in defensive and exploratory contexts. Behavioral Ecology and Sociobiology 76:132.

Clark RW, Bakken GS, Reed EJ, Soni A, 2022. Pit viper thermography: the pit organ used by crotaline snakes to detect thermal contrast has poor spatial resolution. Journal of Experimental Biology 225, jeb2444788. doi:10.1242/jeb.244478

Maag DW, Clark RW, 2022. Safety in coils: predation rates of ambush hunting rattlesnakes are extremely low. Amphibia-Reptilia 43:425-430.

Signore E, Clark RW, Schraft HA, 2022. Temperature-based ambush site selection in sidewinder rattlesnakes (Crotalus cerastes). The Southwestern Naturalist, 65:282-287.

Freymiller GA, Whitford MD, Schwaner MJ, McGowan CP, Higham, TE, Clark RW, 2022. Comparative analysis of Dipodomys species indicates that kangaroo rat hindlimb anatomy is adapted for rapid evasive leaping. Journal of Anatomy 240:466–474. 

Whitford MD,Freymiller GA, Higham, TE, Clark RW, 2022. Shaking things up: the unique feeding behaviour of western banded geckos when consuming scorpions. Biological Journal of the Linnean Society. 2022, doi: 10.1093/biolinnean/blab167

Plein J, Clark RW, Arndt KA, Oechel WC, Stowe D, Zona D, 2022. Response of vegetation and carbon fluxes to brown lemming herbivory in Northern Alaska. Biogeosciences 19, 2779–2794. Doi: 10.5194/bg-2021-286


Teshera ME, Clark, RW, 2021. Strike-induced chemosensory searching in reptiles: a review. Herpetological Monographs 35:28-52. 

Liao CP, Hsu JY, Huang SP, Clark RW, Lin JW, Tseng HY, Huang WS, 2021. Sum of fears among intraguild predators drives the survival of green sea turtle (Chelonia mydas) eggs. Proceedings of the Royal Society B, 288:20202631.

Teshera MS, Clark RW, Wagler AE, Greenbaum E, 2021. Foraging and scavenging behaviour of the prairie rattlesnake (Crotalus viridis): no evidence that envenomation cues facilitate kleptoparasitism of struck prey. Amphibia-Reptilia 42:491-501.

Freymiller GA, Whitford MD, Schwaner MJ, McGowan CP, Higham, TE, Clark RW, 2021. Comparative analysis of Dipodomys species indicates that kangaroo rat hindlimb anatomy is adapted for rapid evasive leaping. Journal of Anatomy 2021, doi: 10.1111/joa.13567 

Robinson KE, Holding ML, *Whitford MD, Saviola A, Yates JR, Clark RW, 2021. Phenotypic and functional variation in venom and venom resistance of two sympatric rattlesnakes and their prey. Journal of Evolutionary Biology, online first. 

Schwaner MJ, *Freymiller GA, Clark RW, McGowan CP, 2021. How to stick the landing: kangaroo rats use their tails to reorient during evasive jumps away from predators. Integrative and Comparative Biology doi:10.1093/icb/icab043 


Hammond JE, Witkowski S, Wison T, Zouvi CA, Goetz NL, Eck N, Clark RW, 2020. Know thine enemy: predator identity influences the response of western banded geckos (Coleonyx variegatus) to chemosensory cues. Journal of Herpetology, 54:480-484.

Whitford MD, Freymiller GA, Higham, TE, Clark RW, 2020. The effects of temperature on the kinematics of rattlesnake predatory strikes in both captive and field environments. Integrative Organismal Biology 2:cov011-13.

Glaudas X, Rice SE, Clark RW, Graham A, 2020. Male energy reserves, mate‐searching activities, and reproductive success: alternative resource use strategies in a presumed capital breeder. Oecologia, 194:415-425.

Levine BA, Schuett GW, Clark RW, Repp RA, Herrmann HW, Booth W, 2020. No evidence of male-biased sexual selection in a snake with conventional Darwinian sex roles.  Royal Society Open Science 7:201261.

Holding MA, Putman BJ, Kong L, Smith JE, Clark RW, 2020. Physiological stress integrates resistance to rattlesnake venom and the onset of risky foraging in California ground squirrels. Toxins 12:617.

Whitford MD, Freymiller GA, Higham, TE, Clark RW, 2020. The effects of temperature on the defensive strike of rattlesnakes. Journal of Experimental Biology 223:jeb223859

Stanley TR, Clark RW, Fisher RN, Rochester CJ, Root SA, Lombardo KJ, Ostermann-Kelm SD, 2020. Long-term monitoring reveals changes in abundance and body size among vertebrate species occupying an insular urban habitat reserve. Conservation Science and Practice 2:734. Co-senior authors.

Ko YW, Liao CP, Clark RW, Hsu JY, Tseng HY, Huang WS, 2020. Aposematic coloration enhances memory ability in an Agamid lizard. Animal Behaviour 161:1-13.

Glaudas X, Rice SE, Clark RW, Graham A, 2020. The intensity of sexual selection, body size and reproductive success in a mating system with male-male combat: Is bigger better? Oikos 129:998-1011.


Whitford MD, Freymiller GA, Higham TE, Clark RW, 2019. Determinants of predation success: How to survive an attack from a rattlesnake. Functional Ecology 33:1099-1109.

Schraft HA, Bakken GS, Clark RW, 2019. Infrared-sensing snakes select ambush orientation based on thermal backgrounds. Scientific Reports 9:3950.

Freymiller GA, *Whitford MD, Higham TE, Clark RW, 2019. Escape dynamics of free-ranging desert kangaroo rats (Rodentia: Heteromyidae) evading rattlesnake strikes. Biological Journal of the Linnean Society 127:164-172.

Schraft HA, Clark RW, 2019. Sensory basis of navigation in snakes: the relative importance of eyes and pit organs. Animal Behaviour 147:77-82. Selected as Editor’s Choice.


Bakken GS, Schraft HA, Cattell RW, Tiu DB, Clark RW. 2018. Cooler snakes respond more strongly to infrared stimuli, but we have no idea why. The Journal of Experimental Biology 221:jeb182121–17

Lion KA, Rice SE, Clark RW. 2018. Genetic patterns in fragmented habitats: a case study for two Peromyscus species in southern California. Journal of Mammalogy 99:923–935.

Schraft HA, Goodman C, Clark RW. 2018. Do free-ranging rattlesnakes use thermal cues to evaluate prey? J. Comp. Physiol. A 204:295–303.

Lee CY, Yo SP, Clark RW, Hsu JY, Liao CP, Tseng HY, Huang WS. 2018. The role of different visual characters of weevils signalling aposematism to sympatric lizard predators. Journal of Zoology 306:36–47.


Rice SE, Moustakas E, Nava R, Glaudas X, Alexander GJ, Clark RW, 2017. Characterization of 11 cross-amplifying microsatellite loci for Bitis arietans (Merrem, 1820). Herpetology Notes 10:329-333.

Freymiller GA, Whitford MD, Higham TE, Clark RW, 2017. Recent interactions with snakes enhance escape performance of desert kangaroo rats (Rodentia: Heteromyidae) during simulated attacks. Biological Journal of the Linnean Society, 122:651-660.

Whitford MD, Freymiller GA, Clark RW. 2017, Avoiding the serpent's tooth: predatoreprey interactions between free-ranging sidewinder rattlesnakes and desert kangaroo rats. Animal Behaviour 130:73-78.  

Ayon RW, Putman, BJ, Clark RW, 2017. Recent encounters with rattlesnakes enhance ground squirrel responsiveness to predator cues. Behavioral Ecology and Sociobiology 71:149.

Putman BJ, Clark, RW, 2017. Behavioral thermal tolerances of free-ranging rattlesnakes (Crotalus oreganus) during the summer foraging season. Journal of Thermal Biology 65:8-15.

Whitford MD, Freymiller GA, Ryan JM, Steele DPJ, Tapia CN, Clark RW, 2017, Chionactic occipitalis: hypomelanism. Herpetology Notes 10:411-412

Schraft HA, Clark RW, 2017. Kangaroo rats change temperature when investigating rattlesnake predators. Physiology & Behavior 173:174-178.

Higham TE, Clark RW, Collins CE, Whitford MD, Freymiller GA, 2017. Rattlesnakes are extremely fast and variable when striking at kangaroo rats in nature: three-dimensional high-speed kinematics at night. Sci Rep 7:40412. 


Rice S, Putman BJ, Schraft H, Clark RW. 2016. Crotalus oreganus helleri (Southern Pacific Rattlesnake), loss of rattle style and matrix. Herpetological Review 47:679. 

Kabes LE, Clark RW. 2016. The use of chemical cues by granite night lizards (Xantusia henshawi) to evaluate potential predation risk. Copeia 104:930–941.

Rice SE, Beasley RR, Lance SL, Jones KL, Clark RW. 2016. Development of 24 polymorphic microsatellite markers for the Island Night Lizard (Xantusia riversiana). Conservation Genetics Resources 8:169–196.

Clark RW, Dorr SW, *Whitford MD, *Freymiller GA, Hein SR. 2016. Comparison of anti-snake displays in the sympatric desert rodents Xerospermophilus tereticaudus (round-tailed ground squirrels) and Dipodomys deserti (desert kangaroo rats). J Mammal 97:1709-1717. 

Rice S, Clark RW. 2016. Xantusia riversiana (Island Night Lizard) amelanism. Herpetological Review 47:675.

Clark RW, 2016. The hunting and feeding behavior of wild rattlesnakes. Pp. 91 - 118 in GW Schuett, RS Reiserer, CF Smith, and MJ Feldner (Eds.), The Rattlesnakes of Arizona. Eco Publishing, Rodeo, New Mexico. 

Schuett GW, Clark RW, Repp RA, Amarello M, Greene HW. 2016. Social behavior of rattlesnakes: a shifting paradigm. Pp. 161-242 in GW Schuett, RS Reiserer, CF Smith, and MJ Feldner (Eds.), The Rattlesnakes of Arizona. Eco Publishing, Rodeo, New Mexico. 

Maritz B, Penner J, Martins M, Crnobrnja-Isailović J, Spear S, Alencar LRV, Sigala-Rodriguez J, Messenger K, Clark RW, Soorae P, Luiselli L, Jenkins C, Green HW. 2016. Identifying global priorities for the conservation of vipers. Biological Conservation, 204:94-102

Pike DA, Clark RW, Manica A, Tseng H-Y, Hsu J-Y, Huang W-S. 2016. Surf and turf: predation by egg-eating snakes has led to the evolution of parental care in a terrestrial lizard. Sci. Rep. 10.1038/srep22207

Putman BJ, Barbour MA, Clark RW. 2016. The foraging behavior of free-ranging rattlesnakes (Crotalus oreganus) in California ground squirrel (Otospermophilus beecheyi) colonies. Herpetologica 72:55-63. 

Clark RW, Dorr SW, Whitford MD, Freymiller GA, Putman BJ. 2016. Activity cycles and foraging behaviors of free-ranging sidewinder rattlesnakes (Crotalus cerastes): the ontogeny of hunting in a precocial vertebrate. Zoology, 119:196-206. 


Putman BJ, Clark, RW. 2015. Habitat manipulation in hunting rattlesnakes (Crotalus species). Southwestern Naturalist, 60:374-377. 

Hoss SK, Deutschman DH, Booth W, Clark RW. 2015. Post‐birth separation affects the affiliative behaviour of kin in a pitviper with maternal attendance. Biological Journal of Linnean Society.  116:637-648.

Putman BJ, Clark RW. 2015. Crotalus oreganus (Northern Pacific Rattlesnake): non-rattling tail display. Herpetological Review 46:269–270. 

Putman BJ, Coss RG, Clark RW. 2015. The ontogeny of antipredator behavior: age differences in California ground squirrels (Otospermophilus beecheyi) at multiple stages of rattlesnake encounters. Behav Ecol Sociobiol 69:1447–1457. 

Santana FE, Swaisgood RR, Lemm JM, Fisher RN, Clark RW. 2015. Chilled frogs are hot: hibernation and reproduction of the Endangered mountain yellow-legged frog Rana muscosa. Endang. Species. Res. 27:43–51.

Putman BJ, Clark RW. 2015. The fear of unseen predators: ground squirrel tail flagging in the absence of snakes signals vigilance. Behavioral Ecology 26:185-193. 


Hoss SK, Clark RW. 2014. Mother cottonmouths (Agkistrodon piscivorus) alter their antipredator behavior in the presence of neonates. Ethology 120:933-941. 

Clark RW, Schuett GA, Repp RA, Amarello M, Smith CF, Herrmann HW. 2014. Mating systems, reproductive success, and sexual selection in a secretive species: A case study of the Western Diamond-Backed Rattlesnake, Crotalus atrox. PLoS ONE 9:e90616. 

Rogers LL, Mansfield SA, Hornby K, Hornby S, Debruyn TD, Mize M, Clark RW, Burghardt GM. 2014. Black bear reactions to venomous and non-venomous snakes in Eastern North America. Ethology 120:641-651.

Brennan PLR, Clark RW, Mock DW. 2014. Time to step up: defending basic science and animal behaviour. Animal Behaviour 94:101-105.  


Clark RW, Tangco S, Barbour MA. 2012. Field recordings reveal factors that influence predatory strike success of free-ranging rattlesnakes (Crotalus spp.). Animal Behaviour 84:183-190. 

Barbour MA, Clark RW. 2012. Ground squirrel tail-flag displays alter both predatory strike and ambush site selection behaviours of rattlesnakes. Proceedings of the Royal Society B Biological Sciences, 279:3827-3833. 

Barbour MA, Clark RW. 2012. Diel cycles in chemosensory behavior of free-ranging rattlesnakes lying in wait for prey. Ethology 118:480-488.

Clark RW, Brown WS, Stechert R, Greene HW. 2012. Cryptic sociality in rattlesnakes (Crotalus horridus) kinship analysis. Biology Letters 8:523-525. 


Joshi SS, Johnson R, Rundus A, Clark RW, Barbour M, and Owings DH. 2011. Robotic squirrel models:  study of squirrel-rattlesnake interaction in laboratory and natural settings. IEEE-RAM 18:59-68. 

Clark RW, Ramirez G. 2011. Rosy boas (Lichanura trivirgata) use chemical cues to identify female mice (Mus musculus) with litters of dependent young.  Herpetological Journal 21:187-191. 

Clark RW, Marchand MN, Clifford BJ, Stechert R, Stephens S.  2011. Decline of an isolated timber rattlesnake (Crotalus horridus) population:  Interactions between climate change, disease, and loss of genetic diversity.  Biological Conservation 144:886-891 

Pre 2010

Clark RW, Brown WS, Stechert R, Zamudio KR, 2010. Roads, interrupted dispersal, and genetic diversity in timber rattlesnakes (Crotalus horridus).  Conservation Biology, 24:1059-1069. 

Clark RW, Brown WS, Stechert R, Zamudio, KR, 2008. Integrating individual behavior and landscape genetics: the population structure of timber rattlesnake hibernacula. Molecular Ecology 17:719-730. . Commentary: Shine, R. 2008.

Clark, R. W. 2007. Public information for solitary foragers: timber rattlesnakes use conspecific chemical cues to select ambush sites. Behavioral Ecology 18:487-480.

Clark, R. W. 2006. Post-strike behavior of timber rattlesnakes (Crotalus horridus) during natural predation events. Ethology 112:1089-1094.

Clark, R. W. 2006. Fixed videography to study the predation behavior of an ambush foraging snake, Crotalus horridus. Copeia 2006:181-187. 

Clark, R. W. 2005.  Pursuit-deterrent communication between prey animals and timber rattlesnakes (Crotalus horridus): the response of snakes to harassment displays. Behavioral Ecology and Sociobiology 59:258-261. 

Clark, R. W. 2004 Feeding experience modifies the assessment of ambush sites by the timber rattlesnake, a sit-and-wait predator. Ethology 110, 471-483.

Clark, R. W. 2004 Kin recognition in rattlesnakes. Proceeding of the Royal Society of London Series B Biology Letters 271, S243–S245.   

Clark, R. W. 2004. Timber rattlesnakes (Crotalus horridus) use chemical cues to select ambush sites. Journal of Chemical Ecology 30, 607-617. 

Clark, R. W. 2002. Diet of the timber rattlesnake, Crotalus horridus. Journal of Herpetology 36, 494-499.


Other Publications

Clark RW, 2021. Book Review: The Secret Social Lives of Reptiles. Herpetological Bulletin 158, doi: 10.33256/hb158.4748 

Whitford MD, Freymiller GA, Clark RW, 2018. Rattlesnakes and Kangaroo Rats.  Desert Report, News of the desert from Sierra Club California & Nevada Desert Committee. December 2018.

Whitford MD, Freymiller GA, Clark RW, 2018. Predator-prey interactions between rattlesnakes and kangaroo rats.  Mojave National Preserve Science Newsletter, 2018.

Clark RW. 2015. Book Review:  How Snakes Work: Structure, Function, and Behavior of the World's Snakes. The Quarterly Review of Biology 90:350. 

Rypien KL, Anderson J, Andras J, Clark RW, Gerrish GA, Mandel JT, Riskin DK, 2007. Students unite to create State of the Planet course. Nature 447:775.

Clark, R. W. 2005. Social lives of rattlesnakes. Natural History, March 2005 Issue.

Last Updated:  December 2023