Department of Earth and Biological Sciences

William K. Hayes, PhD, MS

Biographical index

Research index (each account is accompanied by photographs)

  • Sexual Behavior of Galapagos Marine Iguanas
  • Behavioral Ecology and Function of Sleep in Iguanas
  • Behavioral Ecology and Conservation of Bahamian Seabirds
  • Taxonomy of Potentially Endangered Bahamian Birds
  • Ecology and Taxonomy of Large-billed Savannah Sparrows
  • Behavioral Ecology of Desert Tortoises

Miscellaneous

Why this web site?

This web site is not just about me. Here, my students and I have put together some of the highlights of our various studies. We are linked together because we share a boundless fascination with the natural world. However, the most interesting material showcases not us or our accomplishments, but rather the cherished subjects that we study.

We find it both challenging and rewarding to work with a diversity of projects, but there are several criteria we consider before taking one on. Most important, we choose projects that stimulate our interest (research must be fun) and that promise to provide important new insights (research must be justified). Whether our findings offer useful information to a physician, or simply increase awareness of and appreciation for an organism or threatened environment, we are doing our small part in making this world a better, more interesting place to live in.

We invite you to explore this site and hope that you will enjoy it! Be sure to click on the links to the research accounts, as they offer considerable information about our studies and lots of pictures and videos to browse. Consider visiting the student links as well, where PDFs of theses and dissertations can be downloaded. If you are here to learn about rattlesnake research, there is plenty of good information, but you will also want to visit my personal website, which hosts several informative pages. These include our Biology of the Rattlesnakes Symposium web site, which you can visit to glean abstracts from the many dozens of scientific presentations or learn more about the resulting volume, The Biology of Rattlesnakes, which includes 50 original contributions from 98 authors. You can also learn about the Loma Linda Rattlesnake Study. And if you wish to become better acquainted with The Bahamas, you can also visit the San Salvador's Living Jewels Foundation web site (which is still under development).

If you are contemplating the possibility of working in my lab as a graduate student, please feel free to email me. Bear in mind, however, that my lab is perpetually full and I can anticipate openings only when I have a good idea as to who will graduate at the end of the academic year (I usually know this by January or February). I prefer to work with students who get excited about their projects and are motivated to publish. My students often collaborate with each other, thereby enriching their experience and productivity. Although our program accepts and very much appreciates students from a broad range of backgrounds and beliefs, you should be aware that Loma Linda University is a privately-owned Christian institution. With our emphasis on ecology and conservation, we believe that science and religion--two of the most powerful forces in the world--should work amicably together to save the planet and its fragile ecosystems.

Education

1991 PhD, zoology and physiology, University of Wyoming
1986 MS, biology, Walla Walla College, Washington
1984 BS, biology, Walla Walla College, Washington

Research interests

  • Behavioral ecology of venoms and venomous animals:
    • Venom expenditure by snakes during predatory and defensive bites
    • Venom expenditure by scorpions and centipedes during stinging
    • Snakebite severity and treatment of snakebite
    • The distribution of Mojave toxin among rattlesnake species
    • The biological roles and adaptive value of snake (and other animal) venoms
    • Behavioral ecology of free-ranging southern California rattlesnakes
  • Behavioral ecology and conservation of non-venomous reptiles:
    • Behavioral ecology and conservation of endangered Bahamian rock iguanas
    • Behavioral ecology of Green Iguanas in Costa Rica
    • Lek-mating tactics of Galapagos Marine Iguanas
    • Behavioral ecology and function of sleep in iguanas and other lizards
    • Behavioral ecology of tortoises
  • Behavioral ecology, taxonomy, and conservation of birds:
    • Behavioral ecology and conservation of Bahamian seabirds
    • Conservation taxonomy of Bahamian birds
    • Winter behavioral ecology and taxonomy of Large-billed Savannah Sparrows
    • Behavior and conservation of California Condors

Selected publications

  •  Berube, M. D., S. G. Dunbar, K. Ruetzler, and W. K. Hayes. 2011. Home range and foraging ecology of juvenile Hawksbill Sea Turtles (Eretmochelys imbricata) on inshore reefs of Honduras. Chelonian Conservation and Biology (in press).
  •  Hayes, F. E., and W. K. Hayes. 2011. Seventh-day Adventist faith and environmental stewardship. In H. T. Goodwin (ed.), [book title not yet specified]. Andrews University Press, Berrien Springs, Michigan. PDF preprint (2596 K)
  •  Nisani, Z., and W. K. Hayes. 2011. Defensive stinging by Parabuthus transvaalicus scorpions: risk assessment and venom metering. Animal Behavior (in press). PDF preprint (176 K)
  •  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. PDF reprint (248 K)
  •  Harless, M. L., A. D. Walde, D. K. Delaney, L. L. Pater, and W. K. Hayes. 2010. Sampling considerations for improving home range estimates of Desert Tortoises: effects of estimator, sampling regime, and sex. Herpetological Conservation and Biology 5:374-387. PDF reprint (388 K)
  •  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. PDF reprint (105 K)
  •  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. PDF reprint (239 K)
  •  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. PDF reprint (284 K)
  •  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 “dominica” flavescens). Auk 127:932-939. PDF reprint (1244 K)
  •  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 23:4-18. PDF reprint (1035 K)
  • 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. (731 K)
  •  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. PDF reprint (279 K)
  •  Marlow, 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 Clinical Nutrition 89:1699S-1703S. PDF preprint (67 K)
  •  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. PDF reprint (139 K) 
  • 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. (334 K)
  • 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 PDF reprint (822 K)
  • 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. (761 K)
  • Dugan, E. A., A. Figueroa, and W. K. Hayes. 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. (879 K)
  • 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 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. (886 K)
  • 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. (1048 K)
  • 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. (1494 K)
  • 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. (188 K)
  • 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. (823 K)
  •  Ashton, R., R. Chance, R. Franz, J. D. Groves, J. D. Hardy, Jr., H. S. Harris, Jr., W. Hayes, W. Hildebrand, D. S. Lee, R. Miller, W. S. Sipple, R. Stanley, C. J. Stine, and R. G. Tuck, Jr. 2007. Delusions of science: concerns regarding the unwarranted introduction of Pine Snakes to the Delmarva Peninsula of Maryland. Bulletin of the Maryland Herpetological Society 43:147-158. (183 K)
  • 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. (255 K)
  • Hayes, F. E., and W. K. Hayes. 2007. The great Ivory-billed Woodpecker debate: perceptions of the evidence. Birding 39(2):36-41. (521 K)
  • 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. (593 K)
  • 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. (178 K) [email to request PDF reprint]
  • 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. (100 K)
  • Utt, A. C., N. C. Harvey, W. K. Hayes, and R. L. Carter. 2007. The effects of rearing methods on social behaviors of mentored captive-reared juvenile California Condors. Zoo Biology 27:1-8. (189 K) [email to request PDF reprint]
  • 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. (1480 K)
  • Carter, R. L., W. K. Hayes, V. Voegeli, and S. Voegeli. 2005. Celebrating Biodiversity 2002 - 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. (312 K)
  • 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. American Journal of Emergency Medicine 23:759-762. (82 K)
  • 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. (254 K)
  • Hayes, W. K., and A. W. White. 2005. A record of Cuban/Caribbean Martin (Progne cryptoleuca/dominicensis) for the Bahamas. North American Birds 59:672-673. (611 K)
  • 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. (366 K)
  • 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. Annals of Emergency Medicine 44:599-604. (169 K)
  • 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. (347 K)
  • 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. University of California Press, Berkeley. (450 K)
  • 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. University of California Press, Berkeley. (437 K)
  • 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. University of California Press, Berkeley. (402 K)
  • Hayes, W. K. 2003. Can San Salvador's iguanas and seabirds be saved? Bahamas Journal of Science 11(1):2-8. (420 K)
  • 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. Annals of Emergency Medicine 40:619-624. (69 K)
  • Hayes, W. K., S. S. Herbert, G. C. Rehling, and J. F. Gennaro. 2002. Factors that influence venom expenditure by viperid and other snakes 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. [email to request PDF reprint]
  • 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. Wilderness and Environmental Medicine 11:180-188. (880 K)
  • 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, UK. (109 K)
  • Hayes, W., R. Carter, and N. Goodyear. 2000. Marking techniques and radiotelemetry. Pp. 77-79 in: A. Alberts (ed.), West Indian Iguanas: Status Survey and Conservation Action Plan. IUCN, Gland, UK. (196 K)
  • Hayes, F. E., W. K. Hayes, and T. O. Garnett. 1998. Rediscovery of the Gray-throated Leaftosser (Sclerurus albigularis) on Tobago, West Indies. Caribbean Journal of Science 34(1-2):166-168. (108 K)
  • Hayes, W. K. 1997. Research update: decline of the Sandy Cay Iguana. Iguana Times 6:31. (88 K)
  • Hayes, W. K. 1995. Venom metering by juvenile Prairie Rattlesnakes (Crotalus v. viridis): effects of prey size and experience. Animal Behavior 50:33-40. (99 K)
  • 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. Journal of the International Iguana Society 4:21-30. (1951 K)
  • 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. (351 K)
  • Hayes, W. K., E. A. Verde, and F. E. Hayes. 1994. Cardiac responses during courtship, male-male fighting and other activities in rattlesnakes. Journal of the Tennessee Academy of Science 69:7-9. (406 K)
  • Hayes, W. K. 1993. Effects of hunger on striking, prey-handling, and venom expenditure of Prairie Rattlesnakes (Crotalus v. viridis). Herpetologica 49:305-310. (360 K)
  • 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. Northwestern Naturalist 74:1-9. (493 K)
  • 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. (402 K)
  • 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. Great Basin Naturalist 53:203-206. (213 K)
  • Hayes, W. K. 1992. Prey-handling and envenomation strategies of Prairie Rattlesnakes (Crotalus v. viridis) feeding on mice and sparrows. Journal of Herpetology 26:496-499. (299 K)
  • 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. (380 K)
  • 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. (381 K)
  • Hayes, F. E., W. K. Hayes, K. R. Beaman, and L. E. Harris. 1992. Sleep-like behaviour in the Galapagos Tortoise (Geochelone elephantopus). Herpetology Journal 2:51-53. (250 K)
  • 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. [email to request PDF reprint]
  • Hayes, W. K. 1991. Ontogeny of striking, prey-handling and envenomation behavior of Prairie Rattlesnakes (Crotalus v. viridis). Toxicon 29:867-875. (425 K)
  • Hayes, W. K., and D. Duvall. 1991. A field study of Prairie Rattlesnake predatory strikes. Herpetologica 47:78-81. (380 K)
  • Hayes, W. K. 1991. Envenomation strategies of Prairie Rattlesnakes. PhD Dissertation, University of Wyoming. xi + 156 pp. (2760 K)
  • Duvall, D., D. Chiszar, W. K. Hayes, J. Leonhardt, and M. J. Goode. 1990. Foraging ecology of Prairie Rattlesnakes: laboratory experiments. Journal of Chemical Ecology 16:87-101. (641 K)
  • 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. National Geographic Research 6:457-469. (1165 K)
  • Hayes, F. E. and W. K. Hayes. 1989. Hawk migrations in Beltsville, Maryland. Maryland Birdlife 45:9-11. (126 K)
  • 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. (466 K)
  • Hayes, W. K. 1986. Observations of courtship in the rattlesnake, Crotalus viridis oreganus. Journal of Herpetology 20:246-249. (189 K)
  • 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. (225 K)
  • 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). Bulletin of the Maryland Herpetological Society 20:135-144. (336 K)

Editorial experience

Iguanas Biology and Conservation Book
  • Co-Editor. The Biology of Rattlesnakes. Hayes, W. K., K. R. Beaman, M. D. Cardwell, and S. P. Bush (eds.). 2008. Loma Linda University Press, Loma Linda, California. [See details here]
  • Co-Editor. Proceedings of the 11th Symposium on the Natural History of the Bahamas. Rathcke, B. J., and W. K. Hayes (eds.). 2007. Gerace Research Center, San Salvador, Bahamas. 
  • Co-Editor. Iguanas: Biology and Conservation (peer-reviewed volume). Alberts, A. A., R. L. Carter, W. K. Hayes, and E. P. Martins (eds.). 2004. Univ. Calif. Press, Berkeley. 
  • Editor. Journal of the International Iguana Society / Iguana Times (1994-1996)
  • Co-editor. Bulletin of the Tennessee Valley Herpetological Society (1992-1994)

Conferences, workshops, programs, and special courses organized or taught

  • 2008 Instructor, Desert Snakes: Evolution and Ecology (at Desert Institute, Joshua Tree National Park, California).
  • 2007 Instructor, Venomous Animals of the Desert (at Desert Institute, Joshua Tree National Park, California; October course).
  • 2007 Instructor, Venomous Animals of the Desert (at Desert Institute, Joshua Tree National Park, California; April course).
  • 2007 Instructor, The Biology of Birds (at Winter Wednesday classes, University Seventh-day Adventist Church, Loma Linda, California).
  • 2006 Instructor, Reptiles of the Desert (at Desert Institute, Joshua Tree National Park, California).
  • 2005 Program Co-chairperson (with Beverly Rathcke), Natural History of the Bahamas Symposium (at Gerace Research Center, San Salvador Island, Bahamas)
  • 2005 Instructor, Seabirds Workshop - Institute of Field Ornithology (at Gerace Research Center, San Salvador Island, Bahamas)
  • 2005 Co-organizer, Organizing Committee Chairman, Web Site Creator, Biology of the Rattlesnakes Symposium (at Loma Linda University, Loma Linda, California)
  • 2005 Organizer, San Salvador's Living Jewels conservation education program for San Salvador Island, Bahamas, schoolchildren grades 1-12; supported by grant from Disney Foundation's Conservation Program.
  • 2003 Co-organizer, Celebrating Biodiversity 2002 conservation education program for San Salvador Island, Bahamas, schoolchildren grades 5-11; supported by grant from Disney Foundation's Conservation Program.
  • 1997 Co-organizer, Biosystematics, Behavioral Ecology, and Conservation of Iguanas (annual meetings of the American Society of Ichthyologists and Herpetologists / Herpetologist's League / Society for the Study of Amphibians and Reptiles, at University of Washington, Seattle)
  • 1995 Organizer, Third Annual Conference/Research Expedition of the International Iguana Society (at Bahamian Field Station, San Salvador Island, Bahamas)
  • 1990 Organizer, Colorado-Wyoming Rattlesnake Conference (an informal gathering at University of Wyoming)

Students

I really enjoy working with my graduate students, even more so if I can drop them off on a remote island without any way of getting off until the work is done (just kidding)...

  • Balduff, Naomi (MS, discontinued) - Reproductive behaviors of the endangered Bahamian rock iguana, Cyclura r. rileyi. [Naomi was accepted into and left for a veterinary science program.]
  • Cooper, Allen (MS, in progress) - Behavioral ecology of venom expenditure by Scolopendra centipedes.
  • Corbit, Aaron (MS, in progress) - Rattlesnakes on the edge: behavioral ecology of Red Diamond Rattlesnakes (Crotalus ruber) at the interface with human residential areas. Research website
  • Cyril, Samuel, Jr. (MS, 2001) - "Behavioral ecology of the endangered San Salvador Rock Iguana (Cyclura rileyi rileyi) in the Bahamas." PDF of thesis (327 K)
  • Dugan, Eric A. (PhD, in progress) - Niche partitioning and ecology of two sympatric rattlesnakes in Southern California.
  • Escobar, Ricardo A., III (MS, 2007) - "Post-release success of headstarted Green Iguanas (Iguana iguana) in Costa Rica." PDF of thesis (1670 K)
  • Figueroa, Alex (MS, 2006) - "Radiotelemetry and behavioral ecology of neonate Southern Pacific Rattlesnakes." PDF of thesis (511 K)
  • Fox, Gerad (MS, in progress) - Behavioral ecology of venom expenditure in the Desert Hairy Scorpion (Hadrurus arizonensis).
  • Fry, Shawn K. (MS, 2001) - "Ecology of the endangered Sandy Cay Rock Iguana, Cyclura rileyi cristata, in the Bahamas." PDF of thesis (1352 K)
  • Harless, Meagan (MS, 2007) - "Spatial ecology of the Desert Tortoise: sampling frequency and biological influences." PDF of thesis (1729 K)
  • Herbert, Shelton S. ("Scott") (PhD, 2007) - "Venom expenditure by viperid and elapid snakes: mechanisms, adaptation, and application." PDF of dissertation (399 K)
  • Herbert, Shelton S. ("Scott") (MS, 1998) - "Factors influencing venom expenditure during defensive bites by Cottonmouths (Agkistrodon piscivorus) and Rattlesnakes (Crotalus viridis, Crotalus atrox)." PDF of thesis (321 K)
  • Nelsen, David (MS, in progress) - Behavioral ecology of venom use and venom ontogeny in the Black Widow Spider, (Latrodectus hesperia).
  • Nisani, Zia (PhD, 2008) - "Behavioral and physiological ecology of scorpion venom expenditure: stinging, spraying, and venom regeneration." PDF of dissertation (1550 K)
  • Person, Carl (MS, in progress) - Phylogeography and speciation in select rattlesnake species.
  • Price, Melissa (PhD, in progress) - Taxonomy, behavioral ecology, and conservation of the endangered Bahama Oriole (Icterus dominicensis northropi).
  • Rehling, G. Curtis (MS, 2002) - "Venom expenditure in multiple bites by rattlesnakes and cottonmouths." PDF of thesis (1060 K)
  • Revell, Timothy (PhD, 2005) - "The behavioral ecology of sleep in the Desert Iguana (Dipsosaurus dorsalis)." PDF of dissertation (1880 K)
  • Reynolds, M. Bryant J. (MS, 2006) - "Conservation taxonomy of the Cuban Parrot: plumage, morphology, and flight call variation." PDF of thesis (843 K)
  • Robinette, Valerie (MS, in progress) - Behavioral ecology and conservation of the endangered Bahama Oriole (Icterus dominicensis northropi).
  • Thornton, Benjamin J. (MS, 2000) - "Nesting ecology of the endangered Acklins Bight Rock Iguana, Cyclura rileyi nuchalis, in the Bahamas." PDF of thesis (1852 K) [research conducted under my co-supervision but thesis completed at Andrews University]
  • Trimm, Neville A., Jr. (PhD, 2004) - "Behavioral ecology of Audubon's Shearwaters (Puffinus lherminieri) at San Salvador, Bahamas." PDF of dissertation (3938 K)
  • Trimm, Neville A., Jr. (MS, 2001) - "Ecology of Audubon's Shearwaters (Puffinus lherminieri) at San Salvador, Bahamas." PDF of thesis (923 K)
  • Utt, Amy (PhD, in progress) - Social behaviors and survival of captive-reared juvenile California Condors (Gymnogyps californianus).

Television contributions

It seems inexplicable to me that the most beautiful and irresistible place that I work-in the Bahamas-has yet to catch a producer's attention. Instead, they all want to film the snakes in my lab...or stories related to them. Yeah, snakes are fun and interesting, but...

  • 2008 Evolve: Venom. Produced by and premiered November 2008 on the History Channel. Provided scientific information, a student (Zia Nisani) for on-camera interviews about his own research project, and access to lab facilities.
  • 2007 Rattler: Behind the Fangs. Produced by British Broadcasting Corporation, Bristol, United Kingdom, for National Geographic Channel. Premiered March 2007. Provided scientific information, on-camera interviews, handling of rattlesnakes, students (Eric Dugan, Scott Herbert) for on-camera interviews about their own research projects, access to lab facilities, and a sprinkling of humor. Unfortunately, the segment involving Eric Dugan was cut from the U.S. airing.
  • 2007 World's Worst Venom. Television program produced by Digital Dimensions, Townsville, Queensland, Australia, for National Geographic Channel. Premiered 2007. Provided scientific information, access to students (Scott Herbert, Zia Nisani) and their projects for on-camera interviews and handling of venomous animals (rattlesnakes and arachnids), and access to research facilities. Segments excised from final production.
  • 2004 Venom ER. Ten-part series produced by the British Broadcasting Corporation Wildvision, Bristol, United Kingdom, for Animal Planet. Premiered March 2004. Provided scientific information, on-camera interviews, handling of venomous animals (rattlesnakes and arachnids), students (Eric Dugan, Scott Herbert) for interviews about their own research projects, access to lab facilities, and a segment involving a snakebite to Max, our family dog (who survived the bite).
  • 2002 Venom ER. Original episode produced by the British Broadcasting Corporation Wildvision, Bristol, United Kingdom, for Animal Planet. Premiered August 2002. Provided scientific information, on-camera interviews, handling of rattlesnakes, and access to facilities.
  • 2001 Stings, Fangs, and Spines: Episode IV. Produced by Beyond Productions, Artarmon, NSW, Australia, for Discovery Channel. Premiered Autumn 2001. Provided scientific information, on-camera interviews, and handling of rattlesnakes.
  • 2001 Special Edition: Poison! Produced by National Geographic Television, Washington, DC, for CNBC's National Geographic Explorer. Aired 8 July 2001. Provided a student (Curtis Rehling) and snakes for on-camera interviews and access to facilities.
  • 2001 The United Snakes of America. Produced by National Geographic Television, Washington, DC, for CNBC's National Geographic Explorer. Premiered January 2001. Provided scientific information, on-camera interviews, and handling of rattlesnakes.
  • 2000 Deadly Vipers. Produced by the British Broadcasting Corporation Natural History Unit, Bristol, United Kingdom, for the BBC and Discovery Channel. Premiered Autumn 2000. Provided scientific information and students (Scott Herbert, Curtis Rehling) for on-camera interviews and handling of rattlesnakes.

Filming

Snakebite! A very patient BBC crew films the re-enactment of our family dog, Max's, dreadful snakebite for an episode of Animal Planet's Venom ER. Nicholas seemed to have Max under control, but I'm sure our "acting" otherwise was pretty dismal. Of all the dogs that could have shown up at the vet when the film crew was waiting for a snakebite case, it was really bizarre that ours was the first. After the frightening incident, Max went to a snake "avoidance" class. I think that it has helped, though we reinforce whenever we see a snake in the canyon we love to hike in. Photograph: William K. Hayes.

S. Bush

Sean P. Bush, MD. Dr. Bush is the much-respected envenomation specialist at Loma Linda University Medical Center who stars in Animal Planet's Venom ER. The television producers seek his clinical experience and expertise, but they do find their way to our lab on occasion. Here, Sean is holding "Nero," a Southern Pacific Rattlesnake (Crotalus oreganus helleri) that inflicted a severe bite on a man who experienced unexpected neurotoxic symptoms more typical of a Mojave Rattlesnake (Crotalus scutulatus) bite. I feel fortunate to collaborate with Sean on various projects...and it's reassuring to know that he's in a building only several-hundred meters away in case I'm ever personally in need of his experience and expertise... Photograph: unknown.

Expert witness

I am occasionally asked and sometimes willing to serve as an expert witness. For the several legal cases I have accepted this responsibility, I have provided testimony on the biology, relative danger, and history of domestication of wild reptiles (e.g., iguanas, snakes); the likelihood and risk of snakebite associated with a victim's behavior (i.e., did the victim antagonize the snake? did the victim's story match the circumstances of the bite?); and site inspection for possible presence and relative risk of rattlesnakes. I only accept cases for which I feel comfortable siding with the moral and ethical issues involved.

Why biology? Family secrets revealed...

It's all my dad's fault. At least I think it is. Recent research suggests that the love of nature (or "biophilia" as it has come to be called) has a genetic basis. Considering the facts that my twin brother, Floyd, is a biology professor (just as I am), my older sister, Kathy, is a physician (and was a biology major as an undergraduate), and my younger brother, Rob, is a construction superintendent (um...), the genetic link seems obvious (well, two out of three siblings ain't bad).

My father, Bob, a construction superintendent as well, was a biology major in college who always loved the outdoors. I'm not sure how my mother, Carroll, tolerated it, but when I was young my dad always seemed to have a snake or two in the house. Some of my most cherished memories were of tagging along behind my dad on a "snakehunt," my favorite activity following a weekend picnic. He always seemed to come up with something in the forests and streams of the Maryland piedmont, where my siblings and I grew up. If we were lucky (i.e. begged and cried and carried on long enough), we got to take the snake home. The next morning, I'd show off the snake to my next-door neighbors. And then I'd scream if the darned snake bit me. Of course, it didn't hurt--I faked the reaction just to impress them (right...).

Naturally, things got more serious as my brothers and I got older. There were often up to a dozen snakes in the basement, usually in their cages. As any mother of an ophiodophile can testify, snakes are exceptionally-talented escape artists. Once, my mother ordered us to follow her into our walk-in closet (I don't recall just why), and promptly screamed when she stepped on a large Northern Water Snake. "Oh, thanks, Mom!" one of my quick-thinking (not) brothers exclaimed. "We've been looking for that snake for two weeks now!" She was very pleased to learn this, of course. Just what all moms want to hear. We were very clever at tormenting our mother. We pretty much had the basement all to ourselves.

My parents, no doubt, dreaded every spring. They could always tell when it arrived, because there would be a cacophony of Spring Peepers, Cricket Frogs, and/or Chorus Frogs emanating from our basement menagerie. We had to invent clever means for convincing them that our pets had educational value. They were particularly amused once when we fed seven fireflies to a Green Tree Frog and then turned out the lights. You could see the bugs blinking alright, all in a line that outlined much of the frog's gut. It was awesome. And then the frog died the next day. We didn't think our parents needed to know this latter fact.

Things got more hush-hush when venomous snakes captivated us during high school. For a month, we hid a couple of Florida Pigmy Rattlesnakes (prizes from a SCUBA trip to the Florida Keys) in the camper in the side yard before we figured out a way to conceal them in our bedroom. A month later, while on a family trip to Mississippi, we snuck a Cottonmouth into a suitcase that our unsuspecting dad secured in the car top carrier. Then, after getting our hands on a Timber Rattlesnake, we knew we had to quietly move the hot snakes elsewhere. Things were getting out of hand, and we were growing increasingly concerned about the next cage breakout. Fortunately, a friend at the local college let us move and keep our booty there. A year or so later, when my brother, Floyd, had our snakes on exhibit at a local youth fair, my dad was rather amused to read a label on the Cottonmouth that listed Mississippi as its source. "Mississippi...yeah, right...where'd you get your hands on... No! Don't tell me! Did you really...?" I wish I had been there to see his expression. I hope my mother never reads this.

On occasion, my Dad would ask us, "Why don't you guys get into birds or something else other than herps?" The word "herps," by the way, is slang for herptiles, which includes many things cold, wet, and/or slimy--like lizards, frogs, salamanders, snakes, toads, and a few snake collectors I've met. You wanna get herps? I can tell you how. Anyway, our answer to our father's question was always the same: "Borrrrrring! You can't catch birds and hold them in your hands!" (We actually believed this at the time.) Oddly enough, in seventh grade we drew names from a bowl to see which classmate we would have to buy a Christmas gift for. Floyd and I (being in the same class) drew the names of two of our closest friends. While in a bookstore, we happened upon A Field Guide to Eastern Birds by Roger Tory Peterson, and decided to get each of our friends a copy. The perfect gag gift! We thought this was very funny. But fate as it would be, those two guys drew each of US from the bowl, and they, too, got each of us the SAME stupid book! Go figure! We thought this was very bizarre.

So what should four energetic guys do who are stuck with four identical books on birdwatching? We managed to round up a few binoculars, naturally, and...well...to make a long story short, we made our dad regret his occasional question. We whined and fussed until he drove us to every known birding hotspot within a seventeen-hour drive (or so it seemed) from home. Boy, was he glad when we got our driver's licenses! Floyd, by the way, later authored a book on the birds of Paraguay and coauthored another on the birds of Trinidad and Tobago. Although I am now doing some bird work, my main interest over the years has been herps--mainly snakes.

So you see...all the elements were right there from the start--in the family genes. What more can I say? It's not my fault! And if my web site lags a few years without updating, you can assume that my mom has probably read about the Cottonmouth and other illicit "pets"...

Is animal research acceptable?

There are some who object to research on animals. However, there are two compelling reasons why we disagree with this position.

First, contrary to the inappropriate and irresponsible claims of many animal rights advocates, research on animals really does save human lives. The treatment of snakebite provides a compelling example. Present estimates suggest that, globally, as many as 100,000 snakebite victims die annually. Countless others suffer horrendous morbidity, with gross tissue damage and often-crippling aftereffects. Fortunately, these numbers would be far worse were it not for the one highly effective form of treatment available to us: antivenom. Over the decades, antivenoms to various snake venoms have probably saved hundreds of thousands (perhaps even millions) of lives and have reduced the severity of outcomes for many additional survivors.

But where does antivenom come from? To produce it, snakes must be captured to procure venom. The venom (in minute amounts) must then be injected into another animal, usually a horse or sheep, that serves as an antivenom factory. The immune system of the injected animal produces millions of antibody molecules, which, when purified from the blood, constitute the antivenom. Both procedures can be conducted with minimal stress and no harm to the animals. Because many snakes produce venom having a unique composition, many types of antivenom need to be produced to provide protection against the many species of dangerous snakes. To put it bluntly, there is as yet NO viable alternative to the use of animals to save humans from the ravages of snakebite.

Second, basic natural history research contributes to a wealth of knowledge that greatly influences public attitudes and environmental policy. Without research on animals, for example, how would we learn what species are endangered, what factors are causing their decline, and what habitat requirements are essential for their survival? Without research on animals, how would we recognize the serious impacts that human activities have on the delicate balance of our environment? Without an understanding of animals that only research can provide, who would recognize the need to protect wildlife and sensitive environments, or how to prioritize limited resources for conservation? And who would care about the plight of any animal if the absence of research left us wholly ignorant of its biology?

As important as research on animals may be, we are staunch advocates of animal welfare. Whether the research subjects are eels or eagles, mice or manatees, animals always deserve to be treated well. Today, government regulations and strict institutional oversight ensure humane treatment of animals used in research. At times, we find the process to gain approval of our various projects time-consuming and burdensome, but the alternative--unrestricted animal use--is less attractive. No doubt, some animals would suffer inappropriately and the reputation of those doing legitimate, humane studies would be tarnished. Thus, careful oversight of animal research benefits not only the animals but the research community as well.

Would you like to sponsor our research?

My students and I would be delighted if you or someone you know would like to contribute to one or more of our research endeavors. Becoming a sponsor would offer a number of benefits:

  • Conservation - Contributions support important conservation projects aimed at saving endangered Bahamian species and the sensitive environments they require to survive.
  • Tax-deduction - Any contribution through the University is fully tax-deductible.
  • Education - Contributions support aspiring students seeking professional careers. The level of funding for a student's project can make an enormous difference for his or her future.
  • Leveraged fund-raising - Contributions will encourage others to become sponsors and can be used as matching funds for grants.
  • Visibility - Sponsors will be acknowledged on this web site and in our publications. We would be pleased to consider naming a student fellowship, a specific project, and even the lab itself after a generous donor.

Please click on the DONATE link at the top left or contact me for details on how to make the contribution.