Scientific Publications


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. [rattlesnake_thermal_behavior.pdf]

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

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. [viper_strikes_nature.pdf]

    Press: Covered by dozens of media outlets, including Science Daily, Gizmodo, Daily Mail, Press Enterprise, CNN.

    Videos from this publication viewed over 2 million times, combined across media outlets and YouTube channel.


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. [rattle_loss.pdf]

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. [xantusia_chemical_cues.pdf]

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. [xantusia_primer_note.pdf]

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. [dipodomys_xerospermophilus_antisnake.pdf]

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

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. [rattlesnake_foraging_review.pdf]

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. [rattlesnake_sociality_review.pdf]

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, In press. [viper_conservation_priorities.pdf]

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 [lizard_parental_care.pdf]

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. [oreganus_foraging.pdf]

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. [sidewinder_foraging.pdf]


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

    Press: New Scientist

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. [cottonmouth_kin_maternal.pdf]

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

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.[antipredator_ontogeny.pdf]

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. [chilled_frogs.pdf]

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. [fear_unseen_predators.pdf]


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

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. [atrox_mating_systems.pdf]

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. [bears_snakes.pdf]

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

    Press: Strange Behaviors Blog, Cosmos and Culture Blog


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. [field_snake_strikes.pdf]

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.[tail_flag_displays.pdf]

    Press: Daily Planet interview, KPBS, ScienceNow, MSNBC

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

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

    Press: Huffington Post, MSNBC


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. [robosquirrel.pdf]

    Press:  Time Magazine, MSNBC, Science20, LA Times, CNN

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. [Rosy_boa_chemical_cues.pdf]

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 [NH_snake_decline.pdf]

Pre 2010

Clark RW, Brown WS, Stechert R, Zamudio KR, 2010. Don’t tread on them: roads, interrupted dispersal, and genetic diversity in timber rattlesnakes (Crotalus horridus).  Conservation Biology, 24:1059-1069. [timber_rattlesnakes_roads.pdf]

    Press: Discovery News, Consevation Maven, Cornell Chronicle,

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. [timber_rattlesnake_pop_gen.pdf]
    Commentary: Shine, R. 2008. [shine_commentary.pdf]

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

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

Clark, R. W. 2006. The use of fixed videography in studying the predation behavior of ambush foraging snakes. Copeia 2006:181-187. [fixed_videography.pdf]

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. [pusuit_deterrent.pdf]
    Press: Science News

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. [feeding_experience.pdf]

Clark, R. W. 2004 Kin recognition in rattlesnakes. Proceeding of the Royal Society of London Series B Biology Letters 271, S243–S245. [kin_recognition.pdf]
    Press: National Geographic, BBC, Science News, ScienceNOW, Earthwatch Radio

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

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



Other Publications:

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

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. [Nature_correspondence.pdf]

Clark, R. W. 2005. Social lives of rattlesnakes. Natural History, March 2005 Issue. [Natural_History_Magazine.pdf]