Professor Emeritus

My current research centers around three major themes: evaluation of historical and ecologic determinants of the distributions of tropical organisms (especially amphibians and reptiles); their ecological role in tropical forests; and biogeographic theory.

My most recent publication is a comprehensive handbook on the 385 species of Costa Rican amphibians and reptiles. The book includes keys, diagnostic descriptions, ecological data and a biogeographic analysis of the herpetofauna. The well-known nature photographers Michael and Patricia Fogden provide the book with full color illustrations.


    • A.B., Stanford University, 1950
    • M.A., Stanford University, 1954
    • Ph.D., Stanford University, 1955



    Tropical biology; biogeography; herpetology and evolution.


Awards and Recognitions:

  • 1974-1980 Founder(1963) and President of the Organization for Tropical Studies
  • 1982-2000 Member of the International Commission on Zoological Nomenclature
  • President of the Southern California Academy of Sciences
  • President of the American Society of Ichthyologists and Herpetologists
  • President of the Society of Systematic Biologists
  • 1998 Honorary Member of Academia Nacional de Ciencias de Costa Rica
  • 2000 Henry S. Fitch Award for Excellence in Herpetology
  • Archie F. Carr, Jr. Medal from the Florida Museum of Natural History and the University of Florida
  • 2005 Outstanding Service Award of the American Institute of Biological Sciences
  • 2009 Robert K. Johnson Distinguished Service Award of the American Society of Ichthyologists and Herpetologists

Recent Representative Publications:

    • 2007.
    • Wagler, 1828 and
 Atractus trilineatus 
    • Wagler, 1827 (Reptilia, Serpentes): proposed conservation.
 Bull. Zool. Nomen. 
    • 64(1):68 (with Marinus S. Hoogmoed).

2007. Comment on the proposed conservation of usage of Rana ocellata Linnaeus, 1758 (Amphibia, Anura). Bull. Zool. Nomen. 64(1):68.

2007. Comment on the proposed precedence of Chelodina rugosa Ogilby, 1890 (currently Macrochelodina rugosa; Reptilia Testudines) over Chelodina oblonga Gray, 1841 (Case 3351; see BZN 63:187-193). Bull. Zool. Nomen/ 64(1): 68.

2007. Amphibian and reptile declines over 35 years at La Selva, Costa Rica. Proc Natl. Acad. Sci. USA 104:8352-8356 (with S.M. Whitfield, K. E. Bell, T. Phillippi, M. Sasa, F. BolaZos, G. Chaves, and M.A. Donnelly)

2007. Montane salamanders from the Costa Rica-Panama border region, with description of wo new species of Bolitoglossa.Copeia2007(3):556-565. (with D. B. Wake and J. Hanken).

2007. Dendrobatidae Cope, 1865(1859)(Amphibia, Anura): proposed conservation. Bulletin Zool. Nomen. 64(4):255-260. (with C. W. Myers, D. R. Frost, and T. Grant.)

2007. Three new malodorous rainfrogs of the genus Pristimantis, (Anura: Bracycephalidae) from the Wokomung Massif in west-central Guyana, South America. Zootaxa (1658):39-55. (with D. Bruce Means).

2008. Spectacular new gliding speices of Ecnomiohyla (Anura: Hylidae) from Central Panama. J. Herp. 42(4): 750-759. (with J. R. Medelson, III, E. Griffith, H. Ross, B. Kubicki, and R. Gagliardo).

2009. A checklist of the amphibians and reptiles of Costa Rica: additions and nomenclatural revisions. Zootaxa (2005):1-23. (with F. Bolanos).

Send email to savy1@cox.net