Dissemination of Research Findings

Research offers no benefit to society or to the organisms studied unless it becomes broadly disseminated. The following television appearances and research publications reflect the quality and productivity of our research and our commitment to a high standard of communication. Student names are indicated in bold font.

Television and educational programs

2010 Snakes Underground. National Geographic Channel (William Hayes).
2010 Denim Protects against Rattlesnake Bites (no formal title). Discovery Channel Canada (William Hayes) and student Shelton Herbert).
2009 Snakes. Educational video series, Idyllwild Nature Center, Idyllwild, California (William Hayes).
2008 Evolve: Venom. History Channel (student Zia Nisani).
2007 Rattler: Behind the Fangs. National Geographic Channel (William Hayes and students Eric Dugan and Shelton Herbert).
2007 World's Worst Venom. National Geographic Channel (students Shelton Herbert and Zia Nisani; excised from final production).
2004 Venom ER. Ten-part series, Animal Planet (William Hayes and students Eric Dugan and Shelton Herbert)
2002 Venom ER. Original episode, Animal Planet (William Hayes).
2001 Stings, Fangs, and Spines: Episode IV. Discovery Channel (William Hayes).
2001 Special Edition: Poison! National Geographic Explorer
(student Curtis Rehling).
2001 The United Snakes of America. National Geographic Explorer (William Hayes).
2000 Deadly Vipers. Discovery Channel (students Shelton Herbert and Curtis Rehling).

Volumes edited

2008 The Biology of Rattlesnakes. Hayes, W. K., K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.). Loma Linda University Press, Loma Linda, California.
2007 Proceedings of the 11th Symposium on the Natural History of the Bahamas. Rathcke, B. J., and W. K. Hayes (eds.). Gerace Research Center, San Salvador, Bahamas.
2004 Iguanas: Biology and Conservation (peer-reviewed volume). Alberts, A. A., R. L. Carter, W. K. Hayes, and E. P. Martins (eds.). Univ. Calif. Press, Berkeley.

Publications on behavioral ecology, biodiversity, and conservation of amphibians and reptiles

Babb, R. D., and E. A. Dugan. 2008. Agkistrodon bilineatus (Mexican Cantil). Geographic distribution. Herpetological Review 39:110.
Beaman, K. R., and E. A. Dugan. 2006. Crotalus ruber (Red Diamond Rattlesnake). Catalogue of American Amphibians and Reptiles 840:1-17.
Beaman, K. R., and W. K. Hayes. 2008. Rattlesnakes: research trends and annotated checklist. Pp. 5-16 in W. K. Hayes, K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.), The Biology of Rattlesnakes. Loma Linda University Press, Loma Linda, California.
Cardwell, M. D. 2008. The reproductive ecology of Mohave Rattlesnakes. Journal of Zoology 274:65-76.
Carter, R. L. and W. K. Hayes. 2004. Conservation of an endangered Bahamian iguana. II. Morphological variation and conservation recommendations. Pp. 258-273 in: A. C. Alberts, R. L. Carter, W. K. Hayes, and E. P. Martins (eds.), Iguanas: Biology and Conservation. Univ. Calif. Press, Berkeley.
Carter, R. L., W. K. Hayes, V. Voegeli, and S. Voegeli. 2005. Celebrating Biodiversity 2002 B Conservation education: a responsibility of higher education. Pp. 12-17 in: T. McGrath and S. Buckner (eds.), Proceedings of the 10th Symposium on the Natural History of the Bahamas. Gerace Research Center, San Salvador Island, Bahamas.
Dugan, E. A. 2009. Species accounts for Sceloporus orcutti, Coleonyx switaki, and Petrosaurus mearnsi. Lizards of the Southwest. In: Jones, L. C., and R. E. Lovich (eds.). Rio Nuevo Publishers, Tucson, Arizona.
Dugan, E. A., Y. Cage, and K. Sharrocks. 2008. Tropidodipsas repleta. Geographic distribution. Herpetological Review 39:244 .
Dugan, E. A., and A. Figueroa. 2008. Masticophis mentovarius. Attempted predation and diet. Herpetological Review 39:471.
Dugan, E. A., W. K. Hayes, and A. Figueroa. 2008. Home range size, movements, and mating phenology of sympatric Red Diamond (Crotalus ruber) and Southern Pacific (Crotalus oreganus helleri) Rattlesnakes in southern California. Pp. 353-364 in W. K. Hayes, K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.), The Biology of Rattlesnakes. Loma Linda University Press, Loma Linda, California.
Dugan, E. A., B. Love, and M. G. Figueroa. 2006. Crotalus basiliscus (Mexican West Coast Rattlesnake). Predation. Herpetological Review 37:231.
Dugan, E. A., and S. A. Meyer. 2007. Agkistrodon bilineatus (Mexican Cantil). Reproduction. Herpetological Review 38:85-86.
Duvall, D., D. Chiszar, W. K. Hayes, J. Leonhardt, and M. J. Goode. 1990. Foraging ecology of Prairie Rattlesnakes: laboratory experiments. J. Chem. Ecol. 16:87‑101.
Duvall, D., M. J. Goode, W. K. Hayes, J. K. Leonhardt, and D. Brown. 1990. Prairie Rattlesnake vernal migration: field experimental analyses and survival value in spring. Nat. Geo. Res. 6:457-469.
Escobar, R. A., III, E. Besier, and W. K. Hayes. 2010. Evaluating headstarting as a management tool: post-release success of Green Iguanas (Iguana iguana) in Costa Rica. International Journal of Biodiversity and Conservation 2:204-214.
Figueroa, A., E. A. Dugan, and W. K. Hayes. 2008. Behavioral ecology of neonate Southern Pacific Rattlesnakes (Crotalus oreganus helleri) tracked with externally-attached transmitters. Pp. 365-376 inW. K. Hayes, K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.), The Biology of Rattlesnakes. Loma Linda University Press, Loma Linda, California.
Goldberg, S. R., C. R. Bursey, K. R. Beaman, and E. A. Dugan. 2006. Crotalus basiliscus (Mexican West Coast Rattlesnake). Endoparasites. Herpetological Review 37:94.
Goldberg, S. R., J. Reyes-Velasco, C. R. Bursey, and E. A. Dugan. 2008. Porthidium hespere (Western Hognose Viper). Endoparasites. Herpetological Review 39:99.
Harless, M. L., A. D. Walde, D. K. Delaney, L. L. Pater, and W. K. Hayes. 2009. Home range, spatial overlap, and burrow use of the Desert Tortoise in the west Mojave Desert. Copeia 2009:378-389.
Hayes, F. E., K. R. Beaman, W. K. Hayes, and L. E. Harris, Jr. 1988. Defensive behavior in the Galapagos Tortoise (Geochelone elephantopus), with comments on the evolution of insular gigantism. Herpetologica 44:11‑17.
Hayes, F. E., W. K. Hayes, K. R. Beaman, and L. E. Harris. 1992. Sleep‑like behaviour in the Galapagos Tortoise (Geochelone elephantopus). Herpetol. J. 2:51-53.
Hayes, W. K. 1986. Observations of courtship in the rattlesnake, Crotalus viridis oreganus. J. Herpetol. 20:246-249.
Hayes, W. K. 1997. Research Update: decline of the Sandy Cay Iguana. Iguana Times 6:31.
Hayes, W. K., K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.). 2008. The Biology of Rattlesnakes. Loma Linda University Press, Loma Linda, California.
Hayes, W. K., K. R. Beaman, M. D. Cardwell, and S. P. Bush. 2008. Introduction. Pp. 1-4 in W. K. Hayes, K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.), The Biology of Rattlesnakes. Loma Linda University Press, Loma Linda, California.
Hayes, W., and R. Carter. 2000. Population monitoring. Pp. 79-85 in: A. Alberts (Ed.), West Indian Iguanas: Status Survey and Conservation Action Plan. IUCN, Gland, Switzerland.
Hayes, W. K., and R. L. Carter. 2005. Biology and conservation of Cyclura rileyi, an endangered Bahamian rock iguana. Pp. 69-82 in: T. McGrath and S. Buckner (eds.), Proceedings of the 10th Symposium on the Natural History of the Bahamas. Gerace Research Center, San Salvador Island, Bahamas.
Hayes, W. K., R. L. Carter, S. Cyril, and B. Thornton. 2004. Conservation of an endangered Bahamian iguana. I. Population assessments, habitat restoration, and behavioral ecology. Pp. 232-257 in: A. C. Alberts, R. L. Carter, W. K. Hayes, and E. P. Martins (eds.), Iguanas: Biology and Conservation. Univ. Calif. Press, Berkeley.
Hayes, W., R. Carter, and N. Goodyear. 2000. Marking techniques. Pp. 77-79 in:A. Alberts (Ed.), West Indian Iguanas: Status Survey and Conservation Action Plan. IUCN, Gland, Switzerland.
Hayes, W. K., R. L. Carter, M. Wikelski, and J. A. Sonnentag. 2004. Determinants of lek mating success in male Galapagos Marine Iguanas: behavior, body size, condition, ornamentation, ectoparasite load, and female choice. Pp. 127-147 in: A. C. Alberts, R. L. Carter, W. K. Hayes, and E. P. Martins (eds.), Iguanas: Biology and Conservation. Univ. Calif. Press, Berkeley.
Hayes, W. K., D. Duvall, and G. W. Schuett. 1992. A preliminary report on the courtship behavior of free-ranging Prairie Rattlesnakes, Crotalus v. viridis (Rafinesque), in southcentral Wyoming. Pp. 45-48 in: Strimple, P. D. and J. L. Strimple (eds)., Contributions in Herpetology. Greater Cincinnati Herpetological Society, Cincinnati, Ohio.
Hayes, W. K., D. M. Hayes, D. Brouhard, B. Goodge, and R. L. Carter. 1995. Population status and conservation of the endangered San Salvador Rock Iguana, Cyclura r. rileyi. J. Intl. Iguana Soc. 4:21-30.
Hayes, W. K., E. A. Verde, and F. E. Hayes. 1994. Cardiac responses during courtship, male-male fighting and other activities in rattlesnakes. J. Tenn. Acad. Sci. 69:7-9.
Lovich, R. E., W. Hayes, H. Mushinsky, and G. Rodda. In press. Transect sampling. In M. Foster et al. (eds.), Measuring and Monitoring Biodiversity: Standard Methods for Reptiles. Smithsonian Press, Washington, D.C.
Revell, T. K., and S. G. Dunbar. 2007. The energetic savings of sleep versus temperature in the Desert Iguana, Dipsosaurus dorsalis, at three ecologically relevant temperatures. Comparative Biochemistry and Physiology A 148:393-398.
Revell, T. K., and W. K. Hayes. 2009. Desert Iguanas (Dipsosaurus dorsalis) sleep less when in close proximity to a rattlesnake predator (Crotalus cerastes). Journal of Herpetology 43:29-37.

Publications on behavioral ecology, biodiversity, and conservation of birds

Hayes, F. E. and W. K. Hayes. 1989. Hawk migrations in Beltsville, Maryland. Md. Birdlife 45:9-11.
Hayes, F. E. and W. K. Hayes. 1983. Differential selection of carotenoid pigments in a leucistic Northern Flicker. Md. Birdlife 39:89-90.
Hayes, F. E. and W. K. Hayes. 1983. First Willet in the Maryland piedmont. Md. Birdlife 39:97.
Hayes, F. E., and W. K. Hayes. 2007. The great Ivory-billed Woodpecker debate: perceptions of the evidence. Birding 39:36-41.
Hayes, F. E., W. K. Hayes, and T. O. Garnett. 1998. Rediscovery of the Gray-throated Leaftosser (Sclerurus albigularis) on Tobago, West Indies. Carib. J. Sci. 34(1-2):166-168.
Hayes, W. K. 2003. Can San Salvador=s iguanas and seabirds be saved? Bahamas J. Sci. 11:2-8.
Hayes, W. K. 2006. The urgent need for conservation taxonomy in the Bahamas: new bird species as an example. Bahamas Naturalist and Journal of Science 1:12-24.
Hayes, W. K., R. X. Barry, Z. McKenzie, and P. Barry. 2004. Grand Bahama=s Brown-headed Nuthatch: a distinct and endangered species. Bahamas Journal of Science 12(1):21-28.
Hayes, W. K., E. D. Bracey, M. R. Price, V. Robinette, E. Gren, and C. Stahala. 2010. Population status of the Chuck-will’s-widow (Caprimulgus carolinensis) in the Bahamas. Wilson Journal of Ornithology 122:381-384.
Hayes, W. K., and A. W. White. 2005. A record of Cuban/Caribbean Martin (Progne cryptoleuca/dominicensis) for the Bahamas. North Amer. Birds 59:672‑673.
McKay, B. D., M. B. J. Reynolds, W. K. Hayes, and D. S. Lee. 2010. Evidence supporting the species status of the Bahama Yellow-throated Warbler (Dendroica flavescens). Auk. In press.
Price, M. R., and W. K. Hayes. 2009. Conservation taxonomy of the Greater Antillean Oriole (Icterus dominicensis): diagnosable plumage variation among allopatric populations. Journal of Caribbean Ornithology 22:19-25.
Reynolds, M. B. J., and W. K. Hayes. 2009. Conservation taxonomy of the Cuban Parrot (Amazona leucocephala): variation in morphology and plumage. Journal of Caribbean Ornithology 22:1-18.
Reynolds, M. B. J., W. K. Hayes, and J. W. Wiley. 2010. Geographic variation in the flight call of the Cuban Parrot (Amazona leucocephala) and its taxonomic significance. Journal of Caribbean Ornithology. In press.
Trimm, N. A., Jr., and W. K. Hayes. 2005. Distribution of nesting Audubon=s Shearwaters (Puffinus lherminieri) on San Salvador Island, Bahamas. Pp. 137-145 in: T. McGrath and S. Buckner (eds.), Proceedings of the 10th Symposium on the Natural History of the Bahamas. Gerace Research Center, San Salvador Island, Bahamas.
Utt, A. C., N. C. Harvey, W. K. Hayes, and R. L. Carter. 2008. The effects of rearing methods on social behaviors of mentored captive-reared juvenile California Condors. Zoo Biology 27:1-8.

Publications on behavioral ecology and biodiversity of venom, including human envenomation

Bush, S. P., S. M. Green, T. A. Laack, W. K. Hayes, M. D. Cardwell, and D. A. Tanen. 2004. Pressure immobilization delays mortality and increases intracompartmental pressure after artificial intramuscular rattlesnake envenomation in a porcine model. Ann. Emerg. Med. 44:599-604.
Bush, S. P., S. M. Green, J. A. Moynihan, W. K. Hayes, and M. D. Cardwell. 2002. Crotalidae polyvalent immune Fab (ovine) antivenom is efficacious for envenomations by Southern Pacific Rattlesnakes. Ann. Emerg. Med. 40:619-624.
Bush, S. P., K. G. Hegewald, S. M. Green, M. D. Cardwell, and W. K. Hayes. 2000. Effects of a negative pressure venom extraction device (Extractor) on local tissue injury after artificial rattlesnake envenomation in a porcine model. Wild. Envir. Med. 11:180-188.
Corbett, S. W., B. Anderson, B. Nelson, S. Bush, W. K. Hayes, and M. D. Cardwell. 2005. Most lay people can correctly identify indigenous venomous snakes. Amer. J. Emerg. Med. 23:759-762.
French, W. J., W. K. Hayes, S. P. Bush, M. D. Cardwell, J. O. Bader, and E. Rael. 2004. Mojave toxin in venom of Crotalus helleri (Southern Pacific Rattlesnake): molecular and geographic characterization. Toxicon 44:781-791.
Gennaro, J. F., H. P. Hall, E. R. Casey, and W. K. Hayes. 2007. Neurotropic effects of venoms and other factors that promote prey acquisition. Journal of Experimental Zoology 307A:488-499.
Hayes, W. K. 1991. Ontogeny of striking, prey‑handling and envenomation behavior of Prairie Rattlesnakes (Crotalus v. viridis). Toxicon 29:867-875.
Hayes, W. K. 1992. Prey‑handling and envenomation strategies of Prairie Rattlesnakes (Crotalus v. viridis) feeding on mice and sparrows. J. Herpetol. 26:496-499.
Hayes, W. K. 1992. Factors associated with the mass of venom expended by Prairie Rattlesnakes (Crotalus v. viridis) feeding on mice. Toxicon 30:449-460.
Hayes, W. K. 1993. Effects of hunger on striking, prey-handling, and venom expenditure of Prairie Rattlesnakes (Crotalus v. viridis). Herpetologica 49:305-310.
Hayes, W. K. 1995. Venom metering by juvenile Prairie Rattlesnakes (Crotalus v. viridis): effects of prey size and experience. Anim. Behav. 50:33-40.
Hayes, W. K. 2008. The snake venom-metering controversy: levels of analysis, assumptions, and evidence. Pp. 191-220 in W. K. Hayes, K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.), The Biology of Rattlesnakes. Loma Linda University Press, Loma Linda, California.
Hayes, W. K. and D. Duvall. 1991. A field study of Prairie Rattlesnake predatory strikes. Herpetologica 47:78-81.
Hayes, W. K. and J. G. Galusha. 1984. Effects of rattlesnake (Crotalus viridis oreganus) envenomation upon mobility of male wild and laboratory mice (Mus musculus). Bull. Md. Herpetol. Soc. 20:135-144.
Hayes, W. K., and D. M. Hayes. 1993. Stimuli influencing the release and aim of predatory strikes of the Northern Pacific Rattlesnake, Crotalus v. viridis. Northwest. Nat. 74:1-9.
Hayes, W. K., and F. E. Hayes. 1985. Human envenomation from the bite of the Eastern Garter Snake, Thamnophis s. sirtalis (Serpentes: Colubridae). Toxicon 23:719-721.
Hayes, W. K., and S. S. Herbert. 2010. In reply: Denim, rattlesnakes, and evidence-based scouting (Letter to the Editor by R. J. Hoffman). Annals of Emergency Medicine 56:74-75.
Hayes, W. K., S. S. Herbert, J. R. Harrison, and K. L. Wiley. 2008. Spitting versus biting: differential venom gland contraction regulates venom expenditure in the Black-necked Spitting Cobra, Naja nigricollis nigricollis. Journal of Herpetology 42:453-460.
Hayes, W. K., S. S. Herbert, G. C. Rehling, and J. F. Gennaro. 2002. Factors that influence venom expenditure in viperids and other snake species during predatory and defensive contexts. Pp. 207-233 in: G. W. Schuett, M. Hoggren, M. E. Douglas, and H. W. Greene (eds.), Biology of the Vipers. Eagle Mountain Publ., Eagle Mountain, Utah.
Hayes, W. K., I. I. Kaiser, and D. Duvall. 1992. The mass of venom expended by Prairie Rattlesnakes when feeding on rodent prey. Pp. 383-388 in: Campbell, J. A. and E. D. Brodie, Jr. (eds.), Biology of the Pitvipers. Selva Publ., Tyler, Texas.
Hayes, W. K., P.A. Lavin-Murcio, and K.V. Kardong. 1993. Delivery of Duvernoy's secretion into prey by the Brown Tree Snake, Boiga irregularis (Serpentes: Colubridae). Toxicon 31:881-887.
Hayes, W. K., P. A. Lavin-Murcio, and K. V. Kardong. 1995. Northern Pacific rattlesnakes meter venom when feeding on prey of different sizes. Copeia 1995:337-343.
Hayes, W. K., and S. P. Mackessy. 2010. Sensationalistic journalism and tales of snakebite: are rattlesnakes evolving more toxic venom? Wilderness and Environmental Medicine 21:35-45.
Herbert, S. S., and W. K. Hayes. 2008. Venom expenditure by rattlesnakes and killing effectiveness in rodent prey: do rattlesnakes expend optimal amounts of venom? Pp. 221-228 in W. K. Hayes, K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.), The Biology of Rattlesnakes. Loma Linda University Press, Loma Linda, California.
Herbert, S. S., and W. K. Hayes. 2009. Denim clothing reduces venom expenditure by rattlesnakes striking defensively at model human limbs. Annals of Emergency Medicine 54:830-836.
Jurado, J. D., E. D. Rael, C. S. Lieb, E. Nakayasu, W. K. Hayes, S. P. Bush, and J. A. Ross. 2007. Complement inactivating proteins and intraspecies venom variation in Crotalus oreganus helleri. Toxicon 49:339-350.
Nisani, Z., S. G. Dunbar, and W. K. Hayes. 2007. Cost of venom regeneration in Parabuthus transvaalicus (Arachnida: Buthidae). Comparative Biochemistry and Physiology A 147:509-513.
Nisani, Z., and W. K. Hayes. Under review. Venom squirting behavior of Parabuthus transvaalicus scorpions (Arachnida: Buthidae) serves a defensive role.
Schmidt, D. F., W. K. Hayes, and F. E. Hayes. 1993. The influence of prey movement on the aim of predatory strikes in the rattlesnake, Crotalus viridis. Gr. Basin Nat. 53:203-206.

Publications on general environmental issues

Harlow, H. J., W. K. Hayes, S. Soret, R. L. Carter, E. R. Schwab, and J. Sabate. 2009. Diet and the environment: does it matter? American Journal of Nutrition 89:1699S-1703S.
Hayes, F. E., and W. K. Hayes. In press. Seventh-day Adventist faith and environmental stewardship. In H. T. Goodwin (ed.), [book title not yet specified]. Andrews University Press, Berrien Springs, Michigan.
Rathke, B. J., and W. K. Hayes (eds.). 2007. Proceedings of the 11th Symposium on the Natural History of the Bahamas. Gerace Research Center, San Salvador, The Bahamas.
Rathcke, B. J., and W. K. Hayes. 2007. Preface. Pp. v-vi in B. J. Rathcke and W. K. Hayes (eds.), Proceedings of the 11th Symposium on the Natural History of the Bahamas. 2007. Gerace Research Center, San Salvador, Bahamas.

Student theses and dissertations

Cyril, Samuel, Jr. (MS, 2001) - Behavioral ecology of the endangered San Salvador Rock Iguana (Cyclura rileyi rileyi) in the Bahamas.
Escobar, Ricardo A., III (MS, 2007) - Post-release success of headstarted Green Iguanas (Iguana iguana) in Costa Rica.
Figueroa, Alex (MS, 2006) - Radiotelemetry and behavioral ecology of neonate Southern Pacific Rattlesnakes.
Fry, Shawn K. (MS, 2001) - Ecology of the endangered Sandy Cay Rock Iguana, Cyclura rileyi cristata, in the Bahamas.
Harless, Meagan L. (MS, 2007) - Spatial ecology of the Desert Tortoise: sampling frequency and biological influences.
Herbert, Shelton S. (PhD, 2007) - Venom expenditure by viperid and elapid snakes: mechanisms, adaptation, and application.
Herbert, Shelton S. (MS, 1998) - Factors influencing venom expenditure during defensive bites by Cottonmouths (Agkistrodon piscivorus) and Rattlesnakes (Crotalus viridis, Crotalus atrox).
Nisani, Zia (PhD, 2008) - Behavioral and physiological ecology of scorpion venom expenditure: stinging, spraying, and venom regeneration.
Rehling, G. Curtis (MS, 2002) - Venom expenditure in multiple bites by rattlesnakes and cottonmouths.
Revell, Timothy K. (PhD, 2005) - The behavioral ecology of sleep in the Desert Iguana (Dipsosaurus dorsalis).
Reynolds, M. Bryant J.(MS, 2006) - Conservation taxonomy of the Cuban Parrot: plumage, morphology, and flight call variation.
Thornton, Benjamin J. (MS, 2000) - Nesting ecology of the endangered Acklins Bight Rock Iguana, Cyclura rileyi nuchalis, in the Bahamas.
Trimm, Neville A., Jr. (PhD, 2004) - Behavioral ecology of Audubon's Shearwaters (Puffinus lherminieri) at San Salvador, Bahamas.
Trimm, Neville A., Jr. (MS, 2001) - Ecology of Audubon's Shearwaters (Puffinus lherminieri) at San Salvador, Bahamas.
Utt, Amy C. (PhD, 2010) - Evaluating captive-breeding techniques and reintroduction success of the California Condor (Gymnogyps californianus).