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Phylogenetic Biology of the Burrowing Snake Tribe Sonorini (Colubridae) Item Type text; Electronic Dissertation Authors Holm, Peter, 1959- Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 30/09/2021 03:36:11 Link to Item http://hdl.handle.net/10150/196086 1 PHYLOGENETIC BIOLOGY OF THE BURROWING SNAKE TRIBE SONORINI (COLUBRIDAE) by Peter Alfred Holm ___________________ A Dissertation Submitted to the Faculty of the DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 2 0 0 8 2 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE As members of the Dissertation Committee, we certify that we have read the dissertation prepared by Peter Holm entitled Phylogenetic Biology of the Burrowing Snake Tribe Sonorini (Colubridae) and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of Doctor of Philosophy __________________________________________________________________ Date: December 17, 2007 Judith Bronstein __________________________________________________________________ Date: December 17, 2007 Brian Enquist __________________________________________________________________ Date: December 17, 2007 Peter Reinthal __________________________________________________________________ Date: December 17, 2007 Cecil Schwalbe Final approval and acceptance of this dissertation is contingent upon the candidate’s submission of the final copies of the dissertation to the Graduate College. I hereby certify that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement. _____________________________________________ Date: December 17, 2007 Dissertation Director: Judith Bronstein __________________________________________________________________ Date: December 17, 2007 Co-Chair: Cecil Schwalbe 3 STATEMENT BY AUTHOR This dissertation has been submitted in partial fulfillment of requirements for an advanced degree at The University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the Library. Brief quotations from this dissertation are allowable without special permission, provided that accurate acknowledgment of source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the head of the major department or the Dean of the Graduate College when in his or her judgment the proposed use of the material is in the interests of scholarship. In all other instances, however, permission must be obtained from the author. SIGNED: _______Peter Holm_______________ 4 ACKNOWLEGEMENTS Completion of this project has been a long and eventful journey from my original conception of it in 1994, first presentation of results in La Paz 2000, and to its resurrection and final defense in 2007. This would not have been possible without the assistance and encouragement of many. Charles Lowe welcomed me back into the herpetology lab at the University of Arizona in 1992. He encouraged me to pursue thus project and I am sure he would have liked to do it himself. John Lundberg was willing to take me on as a graduate student even though my interest was in snakes rather than catfishes. I really did enjoy his course on South America. Committee members Judie Bronstein, Alan DeQuiroz, Harry Greene, Phil Hastings, Wayne Maddison, Peter Reinthal, Cecil Schwalbe, and Oscar Ward helped me to overcome many obstacles and provided valuable insight into the science of biology. Graduate coordinators Suzanne LaClair and Sue Whitworth really helped to keep my program alive. I am grateful to Kent Beaman, George Bradley, Mike Douglas, Darrel Frost, Steve Gotte, Harry Greene, Lee Grismer, Michelle Koo, Charles Lowe, John Lundberg, Roy McDiarmid, Robert McCord, Brad Moon, Charlie Painter, Philip Rosen, Greg Schneider, Sally Shelton, John Simmons, Barbara Stein, Thomas Van Devender, Wayne Van Devender, and Humberto Wong for their assistance with loans and workspace. Robert McCord, Thomas Van Devender, and Wayne Van Devender provided their osteological collections. Dennis Cornejo, Brad Moon, Cecil Schwalbe, Thomas Van Devender, and Wayne Van Devender provided many color slides of live specimens. Pete Mayne and Philip Rosen collaborated on field studies. I thank Judie Bronstein, Darrel Frost, David Maddison, Wayne Maddison, Robert McCord, Peter Reinthal, Cecil Schwalbe, and Dale Turner for their comments on one or more versions of the manuscript. I also thank Al Agellon for assistance and patience with my molecular work. The molecular component of this investigation was funded by a grant from the Research Training Group in the Department of Ecology and Evolutionary Biology, University of Arizona. 5 DEDICATION To all those who have gone before me, I owe the world and the promise it holds. To my parents, I owe my interest in science beginning with that first trip to the American Museum of Natural History when I was six. 6 TABLE OF CONTENTS ABSTRACT........................................................................................................................ 9 INTRODUCTION ............................................................................................................ 10 PRESENT STUDY........................................................................................................... 12 APPENDIX A. PART ONE: MORPHOLOGICAL VARIATION AND SPECIATION IN THE BURROWING SNAKE GENUS CHILOMENISCUS (COLUBRIDAE).......... 15 Abstract............................................................................................................................. 16 Introduction....................................................................................................................... 17 Materials and Methods...................................................................................................... 19 Geographic variation......................................................................................................... 20 Taxonomy ......................................................................................................................... 26 Status of unbanded specimens .................................................................................. 27 Probable mislabeled specimens................................................................................ 28 Species Accounts .............................................................................................................. 29 Patterns of Morphological Diversification ....................................................................... 38 Environment.............................................................................................................. 38 Fossorial traits.......................................................................................................... 39 Primitive traits .......................................................................................................... 40 Speciation.......................................................................................................................... 41 Cape Region vicariance............................................................................................ 41 Gulf of California...................................................................................................... 42 Mid-Peninsular seaway ............................................................................................ 43 Reticulate Evolution ......................................................................................................... 44 Unresolved Questions ....................................................................................................... 46 Key to the Species of Chilomeniscus................................................................................ 46 Acknowledgements........................................................................................................... 47 Literature Cited................................................................................................................. 48 Tables, Figures, and Appendices ...................................................................................... 52 APPENDIX B. PART TWO: PHYLOGENETIC SYSTEMATICS OF THE BURROWING SNAKE TRIBE SONORINI (COLUBRIDAE)...................................... 64 7 Abstract............................................................................................................................. 65 Introduction....................................................................................................................... 66 Materials and Methods...................................................................................................... 68 Molecular data set .................................................................................................... 69 Morphological data set............................................................................................. 70 Phylogenetic analysis ..............................................................................................
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