SYSTEMATICS OF THE COMMON GARTER SNAKE, TH A M N O P M S SIRTAUS VO LU M E I A Dissertation Submitted to the graduate faculty of the Louisiana State University and Agricultural and Mechanical College in p a rtia l fu lfillm e n t o f the requirements for the degree of Doctor of Philosophy in The Department of Biological Sciences by John Jeffrey Boundy B. A ., San Jose State U n iversity, 1983 M . A., San Jose State University, 1990 December 1999 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. UMI Number 9960041 UMI* UMI Microform9960041 Copyright 2000 by Bell & Howell Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. Bell & Howell Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGMENTS I wish to thank the members o f my Graduate Committee, through its various permutations: Douglas Rossman (chair), Mark Hafner (co-chair), Robert Chabreck, Michael Fitzsimons, David Good, Dominique Homberger, and Judith Schiebout. I also thank my unofficial committee: Frank Burbrink, Kazuya Naoki, Peggy Sms, and Joseph SlowinskL Loans were received from the following individuals and institutions, listed alphabetically by museum acronym. I wish to thank Beverly Buisson, USFWS Inspector at New Orleans International Airport, for assisting in paperwork needed to return loans to Canada. AM NH American Museum of Natural History, New York: Linda Ford, Darrel Frost ANSP Academy of Natural Sciences, Philadelphia: John Cadle, Ted Daeschler, Edmund Malnate ARK University of Arkansas, Fayetteville: Steven Beaupre ASUMZ Arkansas State University, State University: Stanley Trauth AUM Auburn University Museum, Auburn: Craig Guyer BCB Bryce C. Brown Collection, Baylor University, Waco: David Lintz BYU Brigham Young University, Provo: Jack Sites, Skip Skidmore CA Chicago Academy of Sciences: Mary Hennen CAS California Academy of Sciences, San Francisco: Robert Drewes, L. Henwood, Jens Vindum ChM Charleston Museum, Charleston: Albert Sanders CM Carnegie Museum of Natural History, Pittsburgh: Ellen Censky, Clarence McCoy, Steve Rogers CRC Charles R. Connor Museum, Pullman: Richard Johnson, Kenneth Kardong FM NH Field Museum of Natural History, Chicago: Alan Resetar, Harold Voris ii Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. INHS Illinois Natural History Survey, Champaign; Chris Philips KU University of Kansas Museum of Natural History, Lawrence: William Duellman, Christopher Sheil, John Simmons LACM Los Angeles County Museum, Los Angeles: Robert Bezy, Dave KIzirian, J. Seigel MCZ Museum of Comparative Zoology, Cambridge; John Cadle, Jose Rosado, Leslie Thomas M M NH J.F. Bell Museum, Minneapolis: John Moriarty, Robert Zink MPM Milwaukee Public Museum, Milwaukee: Robert Henderson MSB Museum of Southwestern Biology, Albuquerque: Norman Scott, Howard Snell, Alexandra Snyder MVZ Museum of Vertebrate Zoology, Berkeley: Harry Greene, Barbara Stein, Emily Taylor NCSM North Carolina State Museum, Raleigh: Jeffrey Beane, Alvin Braswell NMC National Museum of Canada, Ottawa: Nancy Boase, Peter Frank, Michael Rankin, Michele Steigerwald * OMNH Oklahoma Museum of Natural History, Norman: Laurie Vitt SMBU Strecker Museum, Baylor University, Waco: Frederick Gehlbach, David Lintz TCWC Texas Cooperative W ildlife Collection, College Station: Lee Fitzgerald, Kathryn Vaughan TNHC Texas Natural History Collection, Austin: David Cannatella UAB University of Alberta, Edmonton: Karl Larsen, W. Roberts UAZ University of Arizona, Tucson: G. Bradley, Charles Lowe UCM University of Colorado Museum, Boulder Alan DeQueiroz, Rosanne Humphrey, M. Leonard, Heather Stein UCSB University of California, Santa Barbara: Marie Holmgren, Samuel Sweet Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. UF Florida State Museum, Gainesville; David Auth UGAMNH University o f Georgia Museum of Natural History, Athens: Elizabeth McGhee UIM University o f Idaho, Moscow; Richard Wallace (a special note of thanks is due Dr. Wallace for coming out of retirement to retrieve and pack the specimens fo r m e). UIM NH University o f Illinois Museum of Natural History, Urbana: John Petzing, Chris Philips UMMZ University o f Michigan Museum of Zoology, Ann Arbor: Arnold Kluge, Ronald Nussbaum, Greg Schneider UMRC University o f Miami Reference Collection, Miami: Julian Lee, Jay Savage UTA University of Texas, Arlington: Jonathan Campbell, Brian Smith UTKVZC University o f Tennessee Vertebrate Zoology Collection, Knoxville: Gordon Burghardt, Alexander Echtemacht Lists of museum holdings were received from the following individuals at their respective institutions: Avila College, Kansas City Robert Powell Burke Museum, University of Washington, Seattle Sievert Rohwer California Polytechnic College, Pomona G lenn Stewart California State University, Fullerton Bayard Brattstrom, W illiam Presch Central Michigan University, M t. Pleasant James Gillingham Clemson University, Clemson Richard Montanucci Cornell University Charles Dardia Creighton University, Omaha James Platz Drake University, Des Moines James Christiansen IV Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Duke University, Durham Joseph B aiky Ernest A. Lina* Collection, Houma Ernie Liner Georgia Southern University, Statesboro David Rostal Highlands Biological Station Richard Bruce Illinois State University, Normal Lauren Brown Indiana State University, Terre Haute Diana Hews Louisiana State University, Shreveport Laurence Hardy Manitoba Museum of Man and Nature, Winnipeg W illiam Preston Marshall University, Huntington Thomas Pauley Miami University Sheldon G uttm an Mississippi Museum of Natural Science Robert Jones Mississippi State University, Mississippi State Ronn A ltig National Museum of Natural History, Washington Ronald Crombie, Kevin DeQueiroz, George Zu g New York State Museum, Albany Joe Bopp, Robert D aniels Northern Illinois University, Dekalb Richard King Oregon State University, Corvallis Joseph Beatty, Douglas Markle Redpath Museum, Montreal David Green Royal Ontario Museum, Toronto Ross M acCulIoch, R obert M urphy Saint Louis University, St. Louis Robert Aldridge San Diego Society of Natural History, San Diego James Berrian, Sally Shelton San Diego State University, San Diego Tod Reeder Savannah River Ecology Laboratory, Aiken W hit Gibbons Seale Mueum, McNeese State Univ., Lake Charles Avery Williams Southeastern Louisiana University Brian Crother, Richard Seigel Southern Illinois University, Carbondale Ronald Brandon Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. State University of New York, Albany Margaret Stewart Sul Ross State University, Alpine James Scudday Tulane Museum of Natural History, Belle Chasse Harold Dundee University of Alabama, Birmingham K en M arion University of California, Riverside Rudolfo Ruibal University of California San Diego, San Diego Gregory Pregill University of Guelph, Guelph James Bogart, Leslie Rye University of Kentucky, Lexington Butch Brodie University of Mississippi, University Edmund Keiser University of Missouri, Columbia Richard Daniel, Raymond Semlitsch University of Nebraska, Lincoln Royce Ballinger, T. Labedz, John Lynch University of Nebraska, Omaha James Fawcett University of New Hampshire, Durham Thomas Kocher University of North Carolina, Asheville James Petranka University of South Florida, Tampa Henry Mushinsky University of The South, Sewanee Nancy Berner University of Texas El Paso, El Paso C a rl Lieb University of Washington, Seattle Sievert Rohwer, other University of Wyoming, Laramie T im Patton Virginia Commonwealth University, Richmond Charles Blem Western Carolina University, Cullowhee Richard Bruce Worcester State College, Worcester Terry Graham The following individuals collected specimens for me: Aaron Bauer, Brack Bartholomew, Joel Carlin, Keith Casdo, Gary Casper, Steve Karsen, Bill Leonard, Armin Meier, Darrell CYQuinn, Douglas Rossman, Steve Shively, Joe Slowinski, and Travis Taggert. v i Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The following individuals provided Information on Thamnophis sirtalis: Ronn Altig, Alvin Braswell, Patrick Gregory, John Jensen, Paul Moler, William Parker, and Chris Woodcock. The following individuals and agencies provided collecting permits: Blue Watson and Bill Vermilion, Louisiana Department of W ildlife Sc fisheries, Randy Trahan, Louisiana Office of State Parks, Tom Mann and Bill Qtusenbeny, Mississippi Department of W ildlife, fisheries and Parks, James Harris and Howard Poitevant, U. S. FishSc W ildlife Service, and John Herce, Washington Department of Fish 4c W ildlife. Thanks are extended to Johnnie Tarver for a work schedule that allowed me to complete my research, and for threatening bodily injury if I didn't stick with the program. Special thanks go to the Lord for creating snakes, to my parents for encouraging my herpetological interests, and to Pearl, John-Paul and Emily Ann for tolerating the inconveniences of a job-family-school triangle. v ii Reproduced with permission of the copyright owner. Further reproduction prohibited without