The Biology of Rattlesnakes

The refereed volume from the 2005 Symposium can be purchased here. A similar volume is planned for the 2011 symposium. Contributing authors can click here for details on how to submit a manuscript.

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Cover of The Biology of Rattlesnakes Volume

Edited by: William K. Hayes, Kent R. Beaman, Michael D. Cardwell,
                 and Sean P. Bush

Foreword by: Gordon W. Schuett

Binding: Cloth with dust jacket

   • 50 chapters
   • 98 authors
   • 606 pages
   • 20 color plates with 92 color images
   • 214 b&w figures and 92 data tables

Publisher: Loma Linda University Press (2008)
ISBN: 978-159410-011-6
Dimensions: approx. 11.2 x 8.8 x 1.8 inches

Price: $105 (regular edition), $225 (signature edition)

Product Description Return to Menu

Due in part to the toxic nature of their venom, rattlesnakes comprise the most popular and well-studied group of snakes in the world. The Biology of Rattlesnakes showcases the finest research to date by investigators encompassing an enormous breadth of expertise. With 50 original contributions from 98 authorities covering a diverse range of topics, this landmark volume will be looked upon as authoritative for years to come. The beautiful, full-color plates depicting many of the more than 30 rattlesnake species adds a tasteful touch.

Reviews (from the back cover) Return to Menu

“Rattlesnakes are foremost among reptiles in eliciting so many emotions of persons coming into contact with them, including fascination, curiosity, awe, and fear. The unique caudal appendage, the rattle, is a remarkable evolutionary experiment that continues to intrigue scientists studying ecology, behavior, and development. This volume provides an admirable compendium and update on all things pertaining to rattlesnakes. In this single tome can be found scholarly treatises of such topics as history of the study of these snakes, morphological and physiological adaptation and variation, behavior and natural history, properties of their venom, and perhaps most importantly, what should and must be done for the conservation of these wonderful creatures.”

Jonathan A. Campbell, Ph.D.
Professor of Biology, University of Texas at Arlington
Co-editor of “Biology of the Pitvipers” and co-author of “The Venomous Reptiles of the Western Hemisphere,” “The Venomous Reptiles of Latin America,” and “Snake Species of the World: A Taxonomic and Geographic Reference”


“A uniquely exhaustive, topically organized presentation of 21st century research on every conceivable aspect of the biology of rattlesnakes, expressed in the words of specialists from the entire field.  It is an advancement of knowledge indispensable not only to those interested in rattlesnakes but also to those concerned with broad topics (e.g., ecology, conservation) of concern to all animal life.”

Hobart M. Smith, Ph.D.
Professor Emeritus of Ecology and Evolutionary Biology, University of Colorado,Boulder
Author of more than 1,000 publications on herpetology, including “Handbook of Lizards” and “Synopsis of the Herpetofauna of Mexico”


The Biology of Rattlesnakes is an update of current research on this biologically and medically important group of venomous snakes. The contributors have been well chosen, the subject matter balanced, and a worthy summary of current work on rattlesnakes well-referenced. This valuable volume continues in the welcome tradition of previous volumes that publish carefully edited contributions arising from open symposia on venomous snakes. This volume is a wonderful and delightful read.”

Kenneth V. Kardong, Ph.D.
Professor of Biology, Washington State University
Author of “An Introduction to Biological Evolution,” “Vertebrates,” “Comparative Anatomy Laboratory Dissection Guide,” and numerous articles on rattlesnake morphology and behavior

About the Editors Return to Menu

William K. Hayes, Ph.D., is a professor of biology in the Department of Earth and Biological Sciences, Loma Linda University, California. He and his students study the behavior, ecology, and conservation of various reptiles and birds, but also focus on the behavioral ecology of venomous animals and their venoms. Their projects include both field and laboratory investigations of rattlesnakes, scorpions, spiders, and centipedes. Professional website.

Kent R. Beaman, M.S., is a research associate in Ichthyology and Herpetology at the Natural History Museum of Los Angeles County, California. He has published articles on a wide range of topics in herpetology. He also coauthored several rattlesnake accounts for the Catalogue of American Amphibians and Reptiles, published by the Society for the Study of Amphibians and Reptiles. Professional website.

Michael D. Cardwell retired as Chief of Specialized Operations from the San Bernardino County, California, Sheriff’s Department in 2004. He recently conducted a multi-year field study of the Mohave Rattlesnake (Crotalus scutulatus). Before leaving southern California to continue his education in biology at the California State University in Sacramento, he contributed to other rattlesnake projects at Loma Linda University. Professional website.

Sean P. Bush, M.D., F.A.C.E.P., is a physician and professor of emergency medicine at the Loma Linda University School of Medicine, Loma Linda, California. As an envenomation specialist, he studies a number of problems associated with venomous bites and stings, including first aid treatment, antivenom safety and efficacy, and factors associated with snakebite severity. His work was the subject of the 10-part television series, Venom ER, aired by Animal Planet. Professional website.

About the Dust Jacket Return to Menu

William Montgomery (Elgin, Texas) produced the stunning artwork on the dust jacket, which portrays a Red Diamond Rattlesnake (Crotalus ruber) in its natural southern California habitat. This large and relatively understudied species abounds in the hills near Loma Linda University, and is the subject of several chapters in this volume. Mr. Montgomery also produced the art used on the dust jackets of Biology of the Pitvipers and Biology of the Vipers. The hand-colored lithograph is available as a limited edition print from the artist (

Table of Contents Return to Menu



Research and History

Rattlesnakes: Research Trends and Annotated Checklist. KENT R. BEAMAN AND WILLIAM K. HAYES

Laurence Monroe Klauber, 1883-1968: Renaissance Man in San Diego. JANET G. KLAUBER

A Curator and His Rattlesnakes: The History of Laurence Monroe Klauber at the San Diego Zoo. KIM LOVICH, CLARK R. MAHRDT, AND KENT R. BEAMAN


Morphological Analysis of the Contact Zone between the Rattlesnakes Crotalus mitchellii stephensi and Crotalus m. pyrrhus. JESSE M. MEIK

Morphology and Physiological Ecology

Functional Specialization of the Extrinsic Venom Gland Musculature within the Crotaline Snakes (Reptilia: Serpentes) and the Role of the M. Pterygoideus Glandulae. BRUCE A. YOUNG AND KATE JACKSON

Geographic Variation in Western Diamondback Rattlesnake (Crotalus atrox) Morphology. CAROL L. SPENCER

Regional Variation and Sexual Dimorphism in Head Form of the Prairie Rattlesnake (Crotalus viridis viridis): Comparisons Using New Analytical Techniques and Collection Methods. MATTHEW T. SMITH AND MICHAEL L. COLLYER

Proximate Determinants of Sexual Size Dimorphism in the Western Diamond-backed Rattlesnake (Crotalus atrox). EMILY N. TAYLOR AND DALE F. DENARDO

Is There an Optimal Length for the Rattlesnake Rattle? BRAD R. MOON AND ALI M. RABATSKY

Annual Variation in Time-Energy Allocation by Timber Rattlesnakes (Crotalus horridus) in Relation to Food Acquisition. STEVEN J. BEAUPRE

Learning and Evolution of Behavior

Learning in Rattlesnakes: Issues and Analysis. CHARLES I. ABRAMSON AND AARON J. PLACE

Caudal Luring as a Precursor in the Evolution of the Rattle: A Test Using an Ancestral Rattlesnake, Sistrurus miliarius barbouri. ALI M. RABATSKY

Behavioral Ecology of Feeding and Defense

Hunting California Ground Squirrels: Constraints and Opportunities for Northern Pacific Rattlesnakes. DONALD H. OWINGS AND RICHARD G. COSS

Ontogeny of Prey Preference in the Southern Pacific Rattlesnake, Crotalus oreganus helleri. JOHN P. LABONTE

Effects of Chemical and Thermal Cues on Striking Behavior and Post-Strike Chemosensory Searching in Rattlesnakes. DAVID CHISZAR AND HOBART M. SMITH

Perspectives on the Regulation of Venom Expulsion in Snakes. BRUCE A. YOUNG

The Snake Venom-Metering Controversy: Levels of Analysis, Assumptions, and Evidence. WILLIAM K. HAYES

Venom Expenditure by Rattlesnakes and Killing Effectiveness in Rodent Prey: Do Rattlesnakes Expend Optimal Amounts of Venom? SHELTON S. HERBERT AND WILLIAM K. HAYES

Factors Influencing the Antipredator Behavior of Mexican Lance-headed Rattlesnakes (Crotalus polystictus) toward Humans. MELISSA AMARELLO, KEVIN BONINE, AND DAVID LAZCANO

Population Ecology and Habitat Use

Sampling Timber Rattlesnakes (Crotalus horridus): Phenology, Growth, Intimidation, Survival, and a Syndrome of Undetermined Origin in a Northern Population. WILLIAM S. BROWN

Annual Variation in Neonate Recruitment in a Florida Population of the Pigmy Rattlesnake, Sistrurus miliarius. TERENCE M. FARRELL, MELISSA A. PILGRIM, AND PETER G. MAY

A Trophic-Based Approach to the Conservation Biology of Rattlesnakes: Linking Landscape Disturbance to Rattlesnake Populations. CHRISTOPHER L. JENKINS AND CHARLES R. PETERSON

Microhabitat Preferences of the Timber Rattlesnake (Crotalus horridus) in the Hardwood Forests of Indiana. SCOTT E. GIBSON, ZACHARY J. WALKER, AND BRUCE A. KINGSBURY

Thermal Ecology

Observations on the Thermal Ecology of Montane Mexican Rattlesnakes. ROBERT W. BRYSON, JR., JAMES M. MUELLER, AND DAVID LAZCANO

Thermal Ecology of Hibernation in a Population of Great Basin Rattlesnakes, Crotalus oreganus lutosus. VINCENT A. COBB AND CHARLES R. PETERSON

Spatial Ecology

Movements, Migrations, and Mechanisms: A Review of Radiotelemetry Studies of Prairie (Crotalus viridis viridis) and Western Rattlesnakes (Crotalus oreganus). DENNIS JØRGENSEN, C. CORMACK GATES, AND DOUGLAS P. WHITESIDE

Coexistence of Rattlesnakes and Military Operations: Occurrence and Spatial Ecology of the Timber Rattlesnake (Crotalus horridus) on the Warren Grove Gunnery Range in the Pinelands of New Jersey. RONALD M. SMITH, WALTER F. BIEN, HAROLD W. AVERY, AND JAMES R. SPOTILA

Seasonal and Annual Variation in Home Range and Movements of Tiger Rattlesnakes (Crotalus tigris) in the Sonoran Desert of Arizona. MATT GOODE, JEFFREY J. SMITH, AND MELISSA AMARELLO

Ecology of the Aruba Island Rattlesnake, Crotalus durissus unicolor. HOWARD K. REINERT, LAURETTA M. BUSHAR, GIAN L. ROCCO, AND R. ANDREW ODUM

Home Range Size, Movements, and Mating Phenology of Sympatric Red Diamond (Crotalus ruber) and Southern Pacific (C. oreganus helleri) Rattlesnakes in Southern California. ERIC A. DUGAN, ALEX FIGUEROA, AND WILLIAM K. HAYES

Behavioral Ecology of Neonate Southern Pacific Rattlesnakes (Crotalus oreganus helleri) Tracked with Externally-Attached Transmitters. ALEX FIGUEROA, ERIC A. DUGAN, AND WILLIAM K. HAYES

Spatial Ecology, Habitat Use, and Survivorship of Resident and Translocated Red Diamond Rattlesnakes (Crotalus ruber). TRACEY K. BROWN, JEFFREY M. LEMM, JEAN-PIERRE MONTAGNE, JEFF A. TRACEY, AND ALLISON C. ALBERTS

Translocation of Venomous Reptiles in the Southwest: A Solution—or Part of the Problem? HUGH K. MCCRYSTAL AND CRAIG S. IVANYI

Reproductive Behavior and Ecology

Reproductive Biology of the Massasauga (Sistrurus catenatus) from South-Central Illinois. ROBERT D. ALDRIDGE, BENJAMIN C. JELLEN, MATTHEW C. ALLENDER, MICHAEL J. DRESLIK, DONALD B. SHEPARD, JASON M. COX, AND CHRISTOPHER A. PHILLIPS

Reproduction in the Eastern Diamond-backed Rattlesnake, Crotalus adamanteus, Under Optimal Conditions in Captivity. NICK CLARK AND FRED ANTONIO

Timber Rattlesnake (Crotalus horridus) Mating Behavior in Southeastern New York: Female Defense in a Search-Based Mating System. EDWIN M. MCGOWAN AND DALE M. MADISON

Morphology, Reproduction, and Habitat Use of a Northern Population of Banded Rock Rattlesnakes, Crotalus lepidus klauberi. DAVID B. PRIVAL

Conservation Ecology and Education

Phenology of Timber Rattlesnakes (Crotalus horridus) in Southern Minnesota: Implications for Conservation. PHILLIP A. COCHRAN

Biological Variation, Management Units, and a Conservation Action Plan for the Timber Rattlesnake (Crotalus horridus). WILLIAM H. MARTIN, WILLIAM S. BROWN, EARL POSSARDT, AND JOHN B. SEALY

Modeling the Landscape Niche Characteristics of Red Diamond Rattlesnakes (Crotalus ruber): Implications for Biology and Conservation. KENNETH J. HALAMA, ADAM J. MALISCH, MICHAEL ASPELL, JOHN T. ROTENBERRY, AND MICHAEL F. ALLEN

Attitudes toward Rattlesnakes by the Peoples of North America and Implications for Rattlesnake Conservation. KIYOSHI SASAKI, AARON J. PLACE, AND KARA N. GAYLOR

The Grass is Rattling: A Rattlesnake Conservation Education Program and Exhibit Made Possible by a Private-Public Partnership. ROBERT L. CARMICHAEL


Venom Composition in Rattlesnakes: Trends and Biological Significance. STEPHEN P. MACKESSY

Phylogeny and the Evolution of ß-Neurotoxic Phospholipases A2 (PLA2) in the Venoms of Rattlesnakes, Crotalus and Sistrurus (Serpentes: Viperidae). STEVEN D. WERMAN

Geographic Distribution of Mojave Toxin and Mojave Toxin Subunits among Selected Crotalus Species. RANDY L. POWELL, CARL S. LIEB, AND EPPIE D. RAEL

Perspective on Venom Evolution in Crotalus. RANDY L. POWELL AND CARL S. LIEB

The Ecological and Evolutionary Context of Mammalian Resistance to Rattlesnake Venoms. JAMES E. BIARDI


Timber Rattlesnake (Crotalus horridus) Envenomations in the Upper Mississippi River Valley. DANIEL E. KEYLER

Important Considerations in Developing Antivenoms. JOHN C. PÉREZ, ROBERT W. FINBERG, AND ELDA E. SÁNCHEZ

Color Plates

Sample Chapters Return to Menu

Rattlesnakes: Research Trends and Annotated Checklist. KENT R. BEAMAN AND WILLIAM K. HAYES  PDF reprint (758 Kb)
<file replaced 3-29-09 with corrected file giving proper common name for Crotalus r. lucasensis>

Hunting California Ground Squirrels: Constraints and Opportunities for Northern Pacific Rattlesnakes. DONALD H. OWINGS AND RICHARD G. COSS  PDF reprint (1.47 Mb)

Seasonal and Annual Variation in Home Range and Movements of Tiger Rattlesnakes (Crotalus tigris) in the Sonoran Desert of Arizona. MATT GOODE, JEFFREY J. SMITH, AND MELISSA AMARELLO  PDF reprint (920 Kb)

Placing orders Return to Menu

Price (regular edition): $105.00
Price (signature edition): $225.00
Price (regular edition, box of 10): $700.00

REGULAR EDITION – The limited-print regular edition includes a dust jacket which the second printing will lack. Supplies will not last long.

SIGNATURE EDITION – The ideal gift for the serious snake lover, researcher, or book collector. The title page includes original black ink signatures from each of the editors and a registered certificate of authenticity. A great value for a great investment.

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