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Lab News:
Shab is graduating.


Al Savitzky holds a snakeAl Savitzky is Department Head and Professor of Biology at Utah State University.  Before moving to USU in July 2011, Al was University Professor of Biological Sciences at Old Dominion University, in Norfolk, VA, where he had served on the faculty since 1982.  Al received his BA from the University of Colorado in 1972 and his MS and PhD from the University of Kansas in 1975 and 1979, respectively.  As a doctoral student Al spent 2 yr as a Smithsonian Predoctoral Fellow, pursuing his dissertation research in the Division of Amphibians and Reptiles, US National Museum of Natural History.  After receiving his doctoral degree Al spent 3.5 yr as a Lecturer at Cornell University.  Al also served two years as a Program Director in the Division of Biological Infrastructure (Human Resource Cluster) at the National Science Foundation.  He also held a three-month appointment as Visiting Professor at the Kyoto University Museum in Japan.  Al has served on the boards of several national and international professional societies and served as President of the Society for the Study of Amphibians and Reptiles and the American Society of Ichthyologists and Herpetologists.  He currently is Treasurer of the World Congress of Herpetology and serves on the Board of Directors of the American Institute for Biological Sciences.

Al’s graduate training was in comparative morphology and morphological systematics, working primarily on the feeding morphology and evolution of colubroid snakes.  He later worked on the embryonic development of snakes, as well as the conservation biology of an endangered rattlesnake, while continuing studies on comparative morphology.  Most recently he has been engaged in a broadly collaborative study of chemical defense  in the Asian snake Rhabdophis tigrinus, which sequesters steroid toxins from toads it consumes as prey and employs those toxin in its own defense.


Al has trained 13 master’s students, eight doctoral students, and three postdoctoral researchers, in addition to his two current doctoral students.

Shab holds a snakeShab Mohammadi  is a doctoral student in Biology at USU, where she is studying the physiological correlates of, and genomic basis for, the tolerance of bufophagous (toad-eating) snakes to bufadienolides toxins.  Shab served for four years as a research assistant at the US National Museum of Natural History, while pursuing her BS degree at George Mason University.  She received her MS degree from Old Dominion University, where she worked under the direction of Al Savitzky, studying the enigmatic enlargement of the adrenal glands in bufophagous snakes.  She began her studies at USU in 2011.  Shab also serves as Curator for the snake pages for the Encyclopedia of Life project. Here's a link to Shab's website.

Akira Mori, Kyoto University

Deborah A. Hutchinson, Coastal Carolina University

Ralph A. Saporito, John Carroll University

Jerrold Meinwald, Cornell University

Frank C. Schroeder, Boyce Thompson Institute for Plant Biology

Victor R. Townsend, Jr., Virginia Wesleyan College

Gordon M. Burghardt, University of Tennessee

John Kleopfer, Virginia Department of Game and Inland Fisheries

Christopher E. Petersen, Naval Facilities Engineering Command

John R. Allsteadt, Virginia Intermont College
PhD student (ODU)
geographic variation in Crotalus horridus

Lisa Behm, Tidewater Community College
MS student (ODU)
behavior of the Aye-aye, Daubentonia madagascariensis

Chad L. Cross, Las Vegas, NV
PhD student (ODU)
movements and habitat use in Agkistrodon piscivorus

Jennifer L. Elwood (deceased)
MS student (ODU)
comparative and developmental morphology of the mandibular symphysis in salamanders

Deborah A. Hutchinson, Coastal Carolina University
MS and PhD student and postdoctoral researcher (ODU)
chemical ecology of Rhabdophis tigrinus

Krista M. McCoy, East Carolina University
postdoctoral researcher (ODU)
adrenal histology of bufophagous snakes; environmental endocrine disrupters in amphibians

Michael J. McCoy, East Carolina University
MS student (ODU)
ecology of Ambystoma mabeei

Konrad Mebert, Zurich, Switzerland
PhD student (ODU)
introgressive hybridization in Nerodia sipedon and N. fasciata

Shabnam Mohammadi, Utah State University
MS student (ODU)
comparative morphology of the adrenal glands in bufophagous snakes

Christopher A. Pague, Colorado Nature Conservancy
PhD student (ODU)
comparative reproductive behavior of anurans

Christopher E. Petersen, Naval Facilities Engineering Command, Atlantic Division
MS student (ODU)
movements and habitat use in Agkistrodon contortrix

Julie A. Ray, LaMICA Biological Station, El Copé, Panama
PhD student (ODU)
ecology of a Panamanian snake community

Ralph A. Saporito, John Carroll University
postdoctoral researcher (ODU)
geographic variation in chemical defense in Rhabdophis tigrinus; chemical ecology of dendrobatid frogs

Robert T. Turner, Virginia Dept. of Environmental Quality
MS student (ODU)
adrenal morphology of Heterodon

William A .Velhagen, Jr., New York University
PhD student (Duke University; co-advised with Louise Roth)
comparative cranial development of natricine snakes

Research in the Savitzky lab generally includes a both field and laboratory approaches and usually includes a morphological component.snakes. Research in the past has included studies of the morphological diversity of feeding mechanisms in snakes, including venom delivery systems and hinged teeth; the embryonic development of the skull of and thermoreceptive pit organs of boas, pythons, and pitvipers; the evolution of the skull in natricine snakes; the anatomy of the tail-shaking muscles of rattlesnakes; and the conservation biology of a suburban population of the Timber Rattlesnake in southeastern Virginia.  Presently the lab is primarily involved in studies of the morphology, physiology, and ecology of sequestered chemical defenses in snakes.
The Yamakagashi is a natricine colubrid snake that possesses a series of defensive structures, known as nuchal glands, in the skin of the neck. The nuchal glands were first described in 1936, and the main component of their secretion was identified in the 1980s as bufadienolide steroids, the same class of toxins that are found in the skin of toads (Bufonidae). Dr. Akira Mori, of the Laboratory of Ethology at Kyoto University (Japan), first suggested that the bufadienolides (BDs) of Rhabdophis may be derived from toads that the snakes consume as prey. In collaboration with Dr. Mori and other colleagues, members of the Savitzky lab demonstrated experimentally that ingested toads are indeed the ultimate source of BDs in Rhabdophis. They further demonstrated that female Rhabdophis can provision their embryos, and thus their newly hatched offspring, with such toxins. These studies are ongoing, and the lab is working with Dr. Savitzky and other colleagues to explore the possible chemical defenses in other members of the Rhabdophis lineage, many of which also possess nuchal glands. We are also pursuing evidence for sequestration in other lineages of bufophagous (toad-eating) snakes, such as Heterodon (Xenodontinae), as well as studying the physiological mechanisms by which bufophagous snakes tolerate the normally lethal effects of BDs on vertebrate heart muscle.
Rhabdophis tigrinus is not the only tetrapod vertebrate known to sequester prey toxins (i.e., store them in its tissues, for use in its own defense). The best-known examples include five independent lineages of so-called “poison frogs,” including the Dendrobatidae of the Neotropics and Mantella in Madagascar, which sequester potent alkaloids from a variety of small arthropod, such as ants and mites. A wealth of circumstantial evidence suggests that many other amphibians and reptiles may also sequester prey toxins, and the subject was recently reviewed by Al Savitzky and colleagues. Our lab is actively seeking students with an interest in pursuing studies in this area (those with a strong background in chemistry are especially encouraged to join the lab). The possibility of sequestered defenses in slug-eating snakes and ant- and mite-eating lizards is of special interest to the lab.
Several studies have been conducted in our lab on the skin of snakes. Gabriel Rivera examined the mechanical properties of snake skin, detailing the degree to which snake skin is adapted to stretch to accommodate intact prey and revealing that snake skin is regionalized along the length of the body. A large comparative study of snake skin, with Drs. Victor R. Townsend, Jr. (Virginia Wesleyan College) and Deborah A. Hutchinson (Coastal Carolina University), has examined the morphological and histological changes in snake skin and its ability to stretch in a sample from basal to more derived taxa.
The Savitzky lab has studied the embryonic development of snakes since the early 1980s, building what is believed to be the largest collection of ophidian (snake) embryos, with about 7,000 specimens of embryos. That collection, curation of which had been supported by the National Science Foundation, was transferred to the Division of Amphibians and Reptiles, U.S. National Museum of Natural History, Smithsonian Institution, in 2011, prior to the lab’s relocation to Utah. Research based on those specimens includes a comparative study of the development of the skull in natricine snakes by William A. Velhagen, Jr., and a study of the development of the skull and thermoreceptive pit organs of pitvipers by Al Savitzky. Ongoing developmental studies include the embryonic development of the nuchal glands in Rhabdophis tigrinus.
The Savitzky lab applies a variety of methods to the study of ophidian structure and function. In addition to gross dissection, the lab makes extensive use of histological methods to examine the microanatomy of tissues such as skin, nuchal glands, adrenal glands, and even entire heads. Scanning electron microscopy is also used on occasion, to examine details of surface structure. We also use vascular casting methods, in which blood vessels are injected with fine latex or resin to reveal details of the arteries, veins, and capillaries associated with various structures.
The Timber Rattlesnake, Crotalus horridus, formerly occurred widely across forested regions of eastern North America, but its populations have been severely reduced and fragmented across much of that range. This has been the result of both intentional killing and, more recently, explosive suburban sprawl into forested areas occupied by rattlesnakes. Our lab conducted studies of movements and habitiat use in two populations in southeastern Virginia from 1982 until about 2008, when the project was taken over fully by two of our primary collaborators, J.D. Kloepfer (Virginia Dept. of Game and Inland Fisheries) and Christopher A. Petersen (Naval Facilities Engineering Command). The longer of the two studies took place on a US Navy base in a forested region of Chesapeake, VA. The study employed radio telemetry to locate individual snakes at intervals usually ranging from daily to monthly, and many were followed for several years. A geographic information system (GIS) was used to analyze the movements of individuals and their use of different habitat types. A subset of this project examined the impact of habitat alteration on the movements of individual snakes. The complete study, based upon well over 20,000 individual location records, is now being prepared for publication.
Savitzky, Alan H., and Joseph T. Collins. 1971. Tantilla gracilis, a snake new to the fauna of Mexico. Journal of Herpetology, 5(1-2):86-87.

Savitzky, Alan H., and Joseph T. Collins. 1971. The ground snake Sonora episcopa episcopa in Coahuila, Mexico. Journal of Herpetology, 5(1-2):87-88.

Savitzky, Alan H., and Hobart M. Smith. 1971. A new snake from Mexico of the taeniata Group of Tantilla. Journal of Herpetology, 5(3-4):167-171.

Cuellar, Hector S., James D. Fawcett, John W. Ferner, Paul Maslin, Jonathan C. Oldham, Jan J. Roth, Alan H. Savitzky, and Hobart M. Smith, 1972. Comment on Dendrobates (Z.N.(S.)1930). Bulletin of Zoological Nomenclature, 29(1):24.

Commins, Martha L., and Alan H. Savitzky. 1973. Field observations on a population of the sand lizard Uma exsul. Journal of Herpetology, 7(1):51-53.

Smith, Hobart M., and Alan H. Savitzky. 1974. Another cryptic associate of the lizard Sceloporus formosus in Guerrero, Mexico. Journal of Herpetology, 8(4):297-303.

Duellman, William E., and Alan H. Savitzky. 1976. Aggressive behavior in a centrolenid frog, with comments on territoriality in anurans. Herpetologica, 32(4):401-404.

Savitzky, Alan H. 1980. The role of venom delivery systems in snake evolution. Evolution, 34(6):1194-1204.

Savitzky, Alan H. 1981. Hinged teeth in snakes: an adaptation for swallowing hard-bodied prey. Science, 212(4492):346-349.

Savitzky, Alan H. 1983. Coadapted character complexes among snakes: fossoriality, piscivory, and durophagy. American Zoologist, 23(2):397-409.

Savitzky, Alan H. 1985. Vertebral protrusion in snakes: Evidence for a novel defensive mechanism. American Philosophical Society Grantees' Reports, 1984:42-43.

Savitzky, Alan H. 1992. Embryonic Development of the Maxillary and Prefrontal Bones of Crotaline Snakes. pp. 119-142. In Campbell, Jonathan A., and E. D. Brodie, Jr., eds. The Biology of Pitvipers. Tyler, TX: Selva.

Buhlmann, Kurt A., Alan H. Savitzky, Barbara A. Savitzky, and Joseph C. Mitchell. 1993. Geographic distribution: Regina rigida. Herpetolological Review, 24(4):156-157.

Velhagen, William A., Jr. and Alan H. Savitzky. 1998. Evolution of embryonic growth in thamnophiine snakes. Copeia, 1998(3):549-558.

Jackson, K., G. Underwood, E. N. Arnold, and Alan H. Savitzky. 1999. Hinged teeth in the enigmatic colubrid, Iguanognathus werneri. Copeia, 1999(3):815-818.

McCoy, Michael W., Donald J. Schwab, and Alan H. Savitzky. 2000. Geographic distribution: Deirochelys reticularia reticularia. Herpetological Review, 31(2):111.

Savitzky, Barbara A., Alan H. Savitzky, Robert T. Belcher, and Scott Ewers. 2002. Geographic distribution: Ramphotyphlops braminus. Herpetological Review, 33(2):150-151.

Hinkley, Jeffrey A., Alan H. Savitzky, Gabriel Rivera, and Stevin H. Gehrke. 2002. Tensile properties of hydrogels and of snake skin. Proceedings of the First World Congress of Biomimetics and Artificial Muscles. Unpaginated. (conference proceedings published as CD-ROM)

Clark, A. M., P. E. Moler, E. E. Possardt, A. H. Savitzky, W. S. Brown, and B. W. Bowen. 2003. Phylogeography of the timber rattlesnake (Crotalus horridus) based on mtDNA sequences. Journal of Herpetology, 37(1):145-154.

Savitzky. Alan H. 2003. Family Colubridae. pp 465-482. In Schlager, Neil, and James B. Murphy eds. Grzimek’s Animal Life Encyclopedia, Revised Edition. Reptiles. Chicago, IL: Gale Group. [invited chapter]

Hutchinson, Deborah A., and Alan H. Savitzky. 2004. Vasculature of the Parotoid Glands of Four Species of Toads (Bufonidae: Bufo). Journal of Morphology, 260(2):247-254.

McCoy, Michael W., and Alan H. Savitzky. 2004. Feeding ecology of larval Ambystoma mabeei. Southeastern Naturalist, 3(3):409-416.

Rivera, Gabriel, Alan H. Savitzky, and Jeffrey A. Hinkley. 2005. Mechanical properties of the integument of the common gartersnake, Thamnophis sirtalis (Serpentes: Colubridae). Journal of Experimental Biology, 208(15):2913-2922.

Allsteadt, John, Alan H. Savitzky, Christopher E. Petersen, and Dayanand N. Naik. 2006. Geographic variation in the morphology of Crotalus horridus (Serpentes: Viperidae). Herpetological Monographs, 20:1-63.

Hutchinson, Deborah A., Akira Mori, Alan H. Savitzky, Gordon M. Burghardt, Xiaogang Wu, Jerrold Meinwald, and Frank C. Schroeder. 2007. Dietary Sequestration of Defensive Steroids in Nuchal Glands of the Asian Snake Rhabdophis tigrinus. Proceedings of the National Academy of Sciences, 104(7):2265-2270.

Montgomery, Chad E., Julie M. Ray, Alan H. Savitzky, Edgardo J. Griffith Rodriguez, Heidi L. Ross, and Karen R. Lips. 2007. Natural History Note: Sibon longifrenis (Drab Snaileater). Diet. Herpetological Review, 38(3):343.

Savitzky, Alan H., and Brad R. Moon. 2008. Tail morphology in the Western Diamond-backed Rattlesnake, Crotalus atrox. Journal of Morphology, 269(8):935-944.

Deborah A. Hutchinson, Alan H. Savitzky, Akira Mori, Jerrold Meinwald, and Frank C. Schroeder. 2008. Maternal provisioning of sequestered defensive steroids by the Asian snake Rhabdophis tigrinus. Chemoecology, 18(3):181-190.

Hutchinson, Deborah A., Alan H. Savitzky, Akira Mori, Gordon M. Burghardt, Jerrold Meinwald, and Frank C. Schroeder. 2011. Investigations of defensive steroid sequestration by the Asian snake Rhabdophis tigrinus. Chemoecology. (DOI: 10.1007/s00049-011-0078-2)

Townsend, Victor R., Jr., Daniel N. Proud, Alan H. Savitzky, and Merry V. Marshall. 2011. Use of coconut as an oviposition site by a Neotropical harvestman (Opiliones, Cranaidae). Living World, 2011:70-71.

Mori, Akira, Gordon M. Burghardt, Alan H. Savitzky, Kathleen A. Roberts, Deborah A. Hutchinson, and Richard C. Goris. Nuchal glands: A novel defensive system in snakes. Chemoecology. (DOI: 10.1007/s00049-011-0086-2)

Savitzky, Alan H., William A. Velhagen, Jr., and Neil Chernoff. 2010. Chapter 6, Appendix B. Collection and Preservation of Reptilian Embryos. In McDiarmid, R. W., M. S. Foster, C. Guyer, J. W. Gibbons, and N. Chernoff (eds.). Reptile Biodiversity: Standard Methods for Inventory and Monitoring. Berkeley: Univ. of California Press.

Ray, Julie M., Chad E. Montgomery, Heidi K. Mahon, Alan H. Savitzky, and Karen R. Lips. 2012 Goo-eaters: Diets of the Neotropical snakes Dipsas and Sibon in central Panama. Copeia, 2012(2):197-202.


Savitzky, Alan H., and Ralph A. Saporito. Sequestration of defensive toxins by tetrapod vertebrates: A symposium in memory of John W. Daly. Dedication. For Chemoecology.

Savitzky, Alan H., Akira Mori, Deborah A. Hutchinson, Ralph A. Saporito, Gordon M. Burghardt, Harvey B. Lillywhite, and Jerrold Meinwald. Sequestered defensive toxins in tetrapod vertebrates: Principles, patterns, and prospects for future studies. For Chemoecology.

Pough, F. Harvey, Robin M. Andrews, John E. Cadle, Martha L. Crump, Alan H. Savitzky, and Kentwood D. Wells. 1998. Herpetology. New York: Prentice Hall, 544 pp. [published October 1997; order of 2nd through 6th authors is alphabetical]

Pough, F. Harvey, Robin M. Andrews, John E. Cadle, Martha L. Crump, Alan H. Savitzky, and Kentwood D. Wells. 2001. Herpetology, Second Edition. New York: Prentice Hall, 612 pp. [published 2000; order of 2nd through 6th authors is alphabetical]

Pough, F. Harvey, Robin M. Andrews, John E. Cadle, Martha L. Crump, Alan H. Savitzky, and Kentwood D. Wells. 2004. Herpetology, Third Edition. New York: Prentice Hall. 726 pp. [published 2003; order of 2nd through 6th authors is alphabetical]
Savitzky, Alan H. 1987. Snakes of the World [review]. Copeia, 1987(1):264-266.

Savitzky, Alan H. 1992. Australian Reptiles & Frogs [review]. Herpetologica, 48(1):144-148.

Savitzky, Alan H. 1997. The Reptiles of Virginia [review]. Copeia, 1997(4): 896-899.

Savitzky, Alan H. 1997. Tales of Giant Snakes: A Historical Natural History of Anacondas and Pythons [review]. Herpetological Natural History, 5(2):187-189. [published in 1998]

Savitzky, Alan H. 2000. Rattlesnake: Portrait of a Predator [review]. Copeia, 2000(2):629-631.

Savitzky, Alan H. 2002. Feeding: Form, Function, and Evolution in Tetrapod Vertebrates [review]. Copeia, 2002(4):1160-1164.

Savitzky, Alan H. 2004. Snakes of the United States and Canada: Natural History and Care in Captivity [review]. Quarterly Review of Biology, 79(4):437.

Savitzky, Alan H. 2005. North American Watersnakes: A Natural History [review]. Quarterly Review of Biology, 80(4):490-491.

Savitzky Lab

Alan H. SavitzkyWelcome to the laboratory of Alan H. Savitzky, in the Department of Biology, Utah State University.  Al Savitzky is a herpetologist, with broad interests in the morphology, ecology, and conservation of amphibians and reptiles, especially snakes.  As an organismal biologist, Al’s interests focus on understanding patterns of evolution among snakes from the perspectives of their anatomy, development, and interactions with prey and predators within natural communities.  Currently, the lab is particularly interested in the topic of sequestered chemical defenses in snakes.


Al Savitzky relocated from Old Dominion University (Norfolk, Virginia), to Utah State University.  Students interested in joining the lab are encouraged to contact Al at

Al and his students interact with faculty and students in the laboratories of Edmund D. Brodie, Jr., and Susannah French.  Other faculty at USU with interests in herpetology include Dean James A. MacMahon, Marty Crump, Karen Mock, and Karen Beard.