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The Coachwhip Snake, Masticophis Flagellum (Shaw): Taxonomy and Di^Trebtjtion

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The Coachwhip Snake, Masticophis Flagellum (Shaw): Taxonomy and Di^Trebtjtion Louisiana State University LSU Digital Commons LSU Historical Dissertations and Theses Graduate School 1968 The oC achwhip Snake, Masticophis Flagellum (Shaw): Taxonomy and Distribution. Larry David Wilson Louisiana State University and Agricultural & Mechanical College Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_disstheses Recommended Citation Wilson, Larry David, "The oC achwhip Snake, Masticophis Flagellum (Shaw): Taxonomy and Distribution." (1968). LSU Historical Dissertations and Theses. 1525. https://digitalcommons.lsu.edu/gradschool_disstheses/1525 This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Historical Dissertations and Theses by an authorized administrator of LSU Digital Commons. For more information, please contact [email protected]. This dissertation has been microfilmed exactly as received 69-4508 WILSON, Larry David, 1940- THE COACHWHIP SNAKE, MASTICOPHIS FLAGELLUM (SHAW): TAXONOMY AND DI^TREBTJTION. Louisiana State University and Agricultural and Mechanical College, Ph.D., 1968 Zoology University Microfilms, Inc., Ann Arbor, Michigan THE COACHWHIP SNAKE, MASTICOPHIS FLAGELLUM (SHAW) TAXONOMY AND DISTRIBUTION A Dissertation Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy in The Department of Zoology fey Larry David Wilson M.S., Louisiana State University, 1965 August, 1968 ACKNOWLEDGMENTS During the course of this study I have been aided by a number of people who have loaned me specimens or who have pro­ vided me with information or advice. First of all, I would like to thank my major professor, Dr. Douglas A. Rossman, who has always encouraged my research on this and other topics and given freely of his advice. I should also like to thank the other members of my committee, Dr. George H. Lowery, Dr. J. Harvey Roberts, Dr. Albert H. Meier, and Dr. Walter J. Harman. To the following people I would like to extend my sincere gratitude for their patience with my requests for the loan of large numbers of a very large snake and/or information or advice: Dr. Steven C. Anderson, Dr. Walter Auffenberg, Dr. Richard J. Baldauf, Dr. Charles M. Bogert, Dr. W. Glen Bradley, Dr. Bryce C. Brown, Dr. E. Milby Burton, Dr. Francis M. Bush, Miss Janalee P. Caldwell, Dr. William J. Cliburn, Mr. Joseph F. Copp, Peter Diekiewicz, Dr. William G. Degenhardt, Dr. James R. Dixon, Dr. Charles L. Douglas, Dr. Neill H. Douglas, Dr. William E. Duellman, Dr. Harold A. Dundee, Dr. B. Gandy, Mr. Glenn Gentry, Dr. Bryan P. Glass, Dr. Harvey L. Gunderson, Mr. Donald E. Hahn, Dr. M. Max Hensley, Mr. M. B. Keasey, III, Dr. Edmund D. Keiser, Mr. C. A. Ketchersid, Dr. John P. Kerr, Dr. Wayne King, Dr. John M. Legler, Dr. Alan E. Leviton, Mr. Ernest A. Liner, Dr. Charles H. Lowe, Jr., Mr. Edmond V. Malnate, Dr. T. Paul Maslin, Dr. ii Clarence J. McCoy, Dr. John S. Mecham, Dr. John D. Miller, Dr. Robert H. Mount, Dr. Michael M. Ovchynnyk, Mr. William M. Palmer, Dr. James A. Peters, Dr. Neil D. Richmond, Dr. Douglas A. Rossman, Mr. Norton M. Rubenstein, Dr. Allan J. Sloan, Dr. Hobart M. Smith, Dr. Henry M. Stephenson, Dr. Wilmer W. Tanner, Dr. Ray C. Telfair, III, Mr. Robert A. Thomas, Dr. Donald W. Tinkle, Dr. Charles F. Walker, Dr. Ernest E. Williams, Mr. Kenneth L. Williams, and Dr. John W. Wright. I would also like to thank the following individuals for contributing information useful in the study: Dr. Andrew Arata, Dr. Ralph W. Axtell, Dr. Joseph R. Bailey, Dr. C. L. Baker, Dr. Rollin H. Baker, Dr. Roger W. Barbour, Dr. Albert P. Blair, Dr. H. T. Boschung, Jr., Mr. James L. Christiansen, Mr. Joseph T. .Collins, Dr. James Dobie, Mr. Arthur C. Echternacht, Dr. Richard Etheridge, Dr. Denzel E. Ferguson, Dr. T. P. Haines, Mr. Laurence M. Hardy, Dr. Robert F. Inger, Dr. James C. List, Mr. Wilfred T. Neill, Dr. Brent B. Nickol, Dr. Kenneth S. Norris, Dr. Gerald G. Raun, Dr. R. J. Robel, Dr. James T. Tanner, and Mr. Richard Thomas. I want to sincerely thank Mr. Howard W. Campbell, Mr. Joseph F. Copp, and my fellow and former fellow graduate students, Mr. Richard M. Blaney, Mr. Donald E. Hahn, Dr. Edmund D. Keiser, Mr. R. Earl Olson, Mr. Gerald C. Schaefer, and Mr. Kenneth L. Williams, for giving freely of their knowledge concerning whipsnakes. I would also like to thank Mr. and Mrs. Roger Conant for the iii use of two photographs from their joint photographic collection and Mr. Richard M. Blaney, Dr. William G. Degenhardt, Dr. Howard K. Gloyd, Mr. Gerald C. Schaefer, and Dr. Robert G. Webb for the use of photographs from their respective collections. I am grateful to Mr. Laurence M. Klauber for the loan of a large amount of data compiled by him during his study of western Masticophis flageHum. My sincere thanks go to Mr. Anthony W. Romano and Dr. Robert M. Stabler for the gift of specimens. Finally, I wish to thank Patricia Blaney who typed large portions of the manuscript and Mr. Richard M. Blaney who is responsible for the reproduction of the majority of the illustra­ tions in this dissertation. TABLE OF CONTENTS PAGE INTRODUCTION ............................................. 1 MATERIALS AND METHODS............... 2 HISTORICAL SUMMARY.......... 5 VALIDITY OF THE GENUS MASTICOPHIS.......................... 7 THE FLAGELLUM GROUP OF MASTICOPHIS ................. 11 COMPOSITION OF THE SPECIES MASTICOPHIS FLAGELLUM ........... 13 SUBSPECIES ACCOUNTS. ................................ 18 MASTICOPHIS FLAGELLUM FLAGELLUM ................... 18 MASTICOPHIS FLAGELLUM TESTACEUS ........................ 36 MASTICOPHIS FLAGELLUM LINEATULUS........................ k9 MASTICOPHIS FLAGELLUM PICEUS............................ 6k MASTICOPHIS FLAGELLUM RUDDOCKI.......................... 76 MASTICOPHIS FLAGELLUM CINGULUM. ...................... 8l MASTICOPHIS FLAGELLUM FULIGINOSUS...................... 102 POPULATIONS OF QUESTIONABLE STATUS...................... 12^ KEY TO THE SUBSPECIES OF ADULT MASTICOPHIS FLAGELLUM ..... 127 SIGNIFICANCE AND EVOLUTION OF COLOR PATTERN . 132 VARIATION IN SCUTELLATION AND PROPORTIONS............... lUl HEAD SCUTELLATION . .................................. lUl BODY SCUTELLATION..................................... 153 PROPORTIONS .................................... 166 HEMIPENIS ..... .................................... 167 ANAL S A C S ............................ 170 v PAGE DENTITION............................................. 170 UMBILICUS............................................. 170 NATURAL HISTORY......................................... 172 FOOD AND FEEDING HABITS..................... 172 REPRODUCTION......... 177 PARASITES AND PREDATORS................................ 179 LOCOMOTION ......................................... 179 DEFENSE.AND ESCAPE .................................... 180 TIME OF ACTIVITY.......................... l8l TEMPERATURE RELATIONSHIPS.................. l8l MISCELLANEOUS BEHAVIOR ................................ 183 DISTRIBUTION AND HABITAT......... 186 FOSSIL HISTORY................................. '199 SUMMARY AND CONCLUSIONS .................................. 20U LITERATURE CITED......... 210 VITA............... 227 LIST OF TABLES NUMBER PAGE 1. Variation in number of supralabials in Masticophis flagelTum.............................. 1^2 2. Variation in number of supralabials enteringorbit in Masticophis flagellum.......................... 1^3 3. Variation in number of infralabials in Masticophis flagellum....... .................... 1^5 I4. Variation in number of infralabials contacting anterior chin shields in Masticophis flagellum .... lb6 5. Variation in number of loreals in Masticophis flagellum.............. 1^7 6. Variation in number of preoculars in Masticophis flagellum.............................. 1^9 7. Variation in number of postoculars in Masticophis flagellum............. 150 8. Percent frequency of deviations observed in the scale reduction pattern of Masticophis flagellum. ........... 168 9. Food of Masticophis flagellum in nature.............. 173 10. Reproductive data for Masticophis flagellum.......... 178 11. Fossil records of Masticophis flagellum. .......... 202 vii LIST OF FIGURES NUMBER PAGE 1. Masticophis flagellum flagellum from 5 miles NW Gainesville, Alachua County, Florida............... 15 2. Distribution of Masticophis flagellum in the United States........... ........................ l6 3. Distribution of Masticophis flagellum in M e x i c o ......... ............................ IT h. Distribution of the light phase of Masticophis flagellum flagellum................................... 23 5. Dorsal view of Masticophis flagellum flagellum, UIMNH 21259, from Key Island, Collier County, Florida........... 26 6. Pattern at midbody of Masticophis flagellum flagellum, CM 2k6k9, from 1.7 miles S Boles, Scott County, Arkansas................ 29 7. Dorsal view of Masticophis flagellum flagellum, NCSM 5U37» from 3 3A miles SSW Ashley Heights, Hoke County, North Carolina........................ 30 8. Masticophis flagellum testaceus, LSUMZ 12878, from Meade County State Lake and Park, Meade County, Kansas ............................... • 3*+ 9. Masticophis flagellum testaceus from the Chisos Mountains, Big Bend National Park, Brewster County, Texas ................ 35 - S ' - 'f 10. Distribution of the phases, of Masticophis flagellum testaceus in the United States ............ 39 11. Percent frequency
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