Here is a list of scientific papers, organized by topic, that have further information on our research topics:


Biardi JE, Chien DC, Coss RG, 2006. California ground squirrel (Spermophilus beecheyi) defenses against rattlesnake venom digestive and hemostatic toxins. Journal of Chemical Ecology 32:137-154.

Coss RG, 1993. Evolutionary persistence of ground squirrel antisnake behavior - reflections. Ecol Psychol 5:171-194

Coss RG, Guse KL, Poran NS, Smith DG, 1993. Development of antisnake defenses in California ground squirrels (Spermophilus beechyi): II. Microevolutionary effects of relaxed selection from rattlesnakes. Behaviour 124:137-164.

Coss RG, Owings DH, 1978. Snake-directed behavior by snake naive and experienced California ground squirrels in a simulated burrow. Zeitschrift Fur Tierpsychologie 48:421-435.

Goldthwaite RO, Coss RG, Owings DH, 1990. Evolutionary dissipation of an antisnake system:  differential behavior by California and arctic ground squirrels in aboveground and belowground contexts. Behaviour 112:246-269.

Hennessy DF, Owings DH, 1978. Snake species discrimination and role of olfactory cues in snake-directed behavior of California ground squirrels. Behaviour 65:115-124.

Hennessy DF, Owings DH, 1988. Rattlesnakes create a context for localizing their search for potential prey. Ethology 77:317-329.

Hennessy DF, Owings DH, Rowe MP, Coss RG, Leger DW, 1981. The information afforded by a variable signal - constraints on snake-elicited tail flagging by California ground squirrels. Behaviour 78:188-226.

Hersek MJ, Owings DH, 1993. Tail flagging by adult California ground squirrels:  a tonic signal that serves different functions for males and females. Animal Behaviour 46:129-138.

Owings DH, Borchert M, Virginia R, 1977. Behavior of California ground squirrels. Animal Behaviour 25:221-230.

Owings DH, Coss RG, 1977. Snake mobbing by California ground squirrels: adaptive variation and ontogeny. Behaviour 62:50-69.

Owings DH, Coss RG, 2008. Hunting California ground squirrels:  constraints and opportuniteis for northern pacific rattlesnakes. In: Biology of the Rattlesnakes (Hayes WK, Beaman KR, Cardwell MD, Bush SP, eds). Loma Linda, CA: Loma Linda University Press.

Owings DH, Coss RG, 2009. Hunting California ground squirrels:  constraints and opportuniteis for northern pacific rattlesnakes. In: Biology of the Rattlesnakes (Hayes WK, Beaman KR, Cardwell MD, Bush SP, eds). Loma Linda, CA: Loma Linda University Press.

Owings DH, Coss RG, McKernon D, Rowe MP, Arrowood PC, 2001. Snake-directed antipredator behavior of rock squirrels (Spermophilus variegatus): Population differences and snake-species discrimination. Behaviour 138:575-595.

Owings DH, Hennessy DF, Leger DW, Gladney AB, 1986. Different functions of alarm calling for different time scales:  a preliminary report on ground squirrels. Behaviour 99:101-116.

Owings DH, Morton ES, 1998. Animal vocal communication:  a new approach. Cambridge: Cambridge University Press.

Owings DH, Owings SC, 1979. Snake-directed behavior by black-tailed prairie dogs (Cynomys ludovicianus). Zeitschrift Fur Tierpsychologie 49:35-54.

Poran NS, Coss RG, 1990. Development of antisnake defenses in California ground squirrels (Spermophilus beecheyi) 1:  behavioral and immunological relationships Behaviour 112:222-245.

Poran NS, Coss RG, Benjamini E, 1987. Resistance of California ground squirrels Spermophilus beecheyi to the venom of the northern pacific rattlesnake Crotalus viridis oreganus:  a study of adaptive variation. Toxicon 25:767-778.

Rundus AS, Owings DH, Joshi SS, Chinn E, Giannini N, 2007. Ground squirrels use an infrared signal to deter rattlesnake predation. PNAS 104:14372-14376.


Bouskila A, 1995. Interactions between predation risk and competition - a field study of kangaroo rats and snakes. Ecology 76:165-178.

Randall JA, 1993. Behavioral adaptations of desert rodents (Heteromyidae). Animal Behaviour 45:263-287.

Randall JA, Hatch SM, Hekkala ER, 1995. Interspecific variation in antipredator behavior in sympatric species of kangaroo rat. Behavioral Ecology and Sociobiology 36:243-250.

Randall JA, King DKB, 2001. Assessment and defence of solitary kangaroo rats under risk of predation by snakes. Animal Behaviour 61:579-587.

Randall JA, Matocq MD, 1997. Why do kangaroo rats (Dipodomys spectabilis) footdrum at snakes? Behavioral Ecology 8:404-413.

Randall JA, Stevens CM, 1987. Footdrumming and other antipredator responses in the bannertail kangaroo rat (Dipodomys spectabalis). Behavioral Ecology and Sociobiology 20:187-194.


Caro TM, 1986. The functions of stotting in Thomson gazelles:  some tests of the predictions. Animal Behaviour 34:663-684.

Caro TM, 1994. Ungulate antipredator behavior:  preliminary and comparative data from African bovids. Behaviour 128:189-228.

Caro TM, 1995. Pursuit-deterrence revisited. Trends in Ecology & Evolution 10:500-503.

Caro TM, 2005. Antipredator defenses in birds and mammals. Chicago: University of Chicago Press.

Clark RW, 2005a. 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.

Fitzgibbon CD, 1989. A cost to individuals with reduced vigilance in groups of Thomsons gazelles hunted by cheetahs. Animal Behaviour 37:508-510.

Fitzgibbon CD, 1994. The costs and benefits of predator inspection behavior in Thomson gazelles. Behavioral Ecology and Sociobiology 34:139-148.

Fitzgibbon CD, Fanshawe JH, 1988. Stotting in Thomson gazelles:  an honest signal of condition. Behavioral Ecology and Sociobiology 23:69-74.

Guilford T, Dawkins MS, 1991. Receiver psychology and the evolution of animal signals. Animal Behaviour 42:1-14.

Hebets EA, Papaj DR, 2005. Complex signal function: developing a framework of testable hypotheses. Behavioral Ecology and Sociobiology 57:197-214.

Laiolo P, Tella JL, Carrete M, Serrano D, Lopez G, 2004. Distress calls may honestly signal bird quality to predators. Proceedings of the Royal Society of London Series B-Biological Sciences 271:S513-S515.

Leal M, 1999. Honest signalling during prey-predator interactions in the lizard Anolis cristatellus. Animal Behaviour 58:521-526.

Leal M, Rodriguezrobles JA, 1995. Antipredator responses of Anolis cristatellus (Sauria, Polychrotidae). Copeia:155-161.

Lima SL, 2002. Putting predators back into behavioral predator-prey interactions. Trends in Ecology & Evolution 17:70-75.

Searcy WA, Nowicki S, 2005. The evolution of animal communication. Princeton, NJ: Princeton University Press.

Seyfarth RM, Cheney DL, 2003. Signalers and receivers in animal communication. Annu Rev Psychol 54:145-173.

Speed MP, 2000. Warning signals, receiver psychology and predator memory. Animal Behaviour 60:269-278.

Zuberbuhler K, Jenny D, Bshary R, 1999. The predator deterrence function of primate alarm calls. Ethology 105:477-490.


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

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

Diller LV, Johnson DR, 1988. Food habits, consumption rates, and predation rates of western rattlesnakes and gopher snakes in southwestern Idaho. Herpetologica 44:228-233.

Duvall D, Chiszar D, Hayes WK, Leonhardt JK, Goode MJ, 1990. Chemical and behavioral ecology of foraging in prairie rattlesnakes. Journal of Chemical Ecology 16:87-102.

Fitch HS, 1949. Study of snake populations in central California. American Midland Naturalist 41:513-579.

Greene HW, 1992. The behavioral and ecological context for pitviper evolution. In: Biology of the Pitvipers (Campbell JA, Brodie Jr. ED, eds). Tyler, TX: Selva; 107-117.

Greene HW, 1997. Snakes:  the evolution of mystery in nature. Berkeley, CA: University of California Press.

Klauber LM, 1972. Rattlesnakes:  their habits, life histories, and influence on mankind. Berkeley: University of California Press.

Reinert HK, Cundall D, Bushar LM, 1984. Foraging behavior of the timber rattlesnake, Crotalus horridus. Copeia 1984:976-981.


Joshi S, Schank J, Giannini N, Hargreaves L, Bish R, 2004. Development of autonomous robotics technology for the study of rat pups. In: Proceedings of the IEEE International Conference on Robotics and Automation. New Orleans, LA; 2860-2864.

Knight J, 2005. When robots go wild. Nature 434:954-955.

Martins El, Ord T, Davenport S, 2005. Combining motions into complex displays: playbacks with a robotic lizard. Behavioral Ecology and Sociobiology 58:351-360.

May CJ, Schank JC, Joshi S, Tran J, Taylor RJ, Scott IE, 2006. Rat pups and random robots generate similar self-organized and intentional behavior. Complexity 12:53-66.

Partan SR, Larco CP, Owens MJ, 2009. Wild tree squirrels respond with multisensory enhancement to conspecific robot alarm behaviour. Animal Behaviour 77:1127-1135.

Patricelli GL, Coleman SW, Borgia G, 2006. Male satin bowerbirds, Ptilonorhynchus violaceus, adjust their display intensity in response to female startling: an experiment with robotic females. Animal Behaviour 71:49-59.

Patricelli GL, Uy JAC, Walsh G, Borgia G, 2002. Sexual selection: Male displays adjusted to female's response. Nature 415:279-280

Schank JC, May CJ, Tran JT, Joshi SS, 2004. A biorobotic investigation of Norway rat pups (Rattus norvegicus) in an arena. Adapt Behav 12:161-173.

Taylor RC, Klein BA, Stein J, Ryan MJ, 2008. Faux frogs: multimodal signalling and the value of robotics in animal behaviour. Animal Behaviour 76:1089-1097.