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Governs the Making of Photocopies Or Other Reproductions of Copyrighted Materials Warning Concerning Copyright Restrictions The Copyright Law of the United States (Title 17, United States Code) governs the making of photocopies or other reproductions of copyrighted materials. Under certain conditions specified in the law, libraries and archives are authorized to furnish a photocopy or other reproduction. One of these specified conditions is that the photocopy or reproduction is not to be used for any purpose other than private study, scholarship, or research. If electronic transmission of reserve material is used for purposes in excess of what constitutes "fair use," that user may be liable for copyright infringement. University of Nevada, Reno Isolation at Work: Body Size Divergence between Reptiles of Nevada’s Pyramid Lake and Anaho Island A thesis submitted in partial fulfillment of the requirements for the degree of Bachelor of Science in Wildlife Ecology and Conservation and the Honors Program by Jade E. Keehn Chris R. Feldman, Ph.D., Thesis Advisor Department of Biology Nathan C. Nieto, Ph.D., Thesis Advisor Department of Agriculture, Nutrition, and Veterinary Sciences May 2012 UNIVERSITY OF NEVADA THE HONORS PROGRAM RENO We recommend that the thesis prepared under our supervision by JADE E KEEHN entitled Isolation at Work: Body Size Divergence between Reptiles of Nevada’s Pyramid Lake and Anaho Island be accepted in partial fulfillment of the requirements for the degree of WILDLIFE ECOLOGY AND CONSERVATION, BACHELOR OF SCIENCE ______________________________________________ Chris R. Feldman, Ph.D., Thesis Advisor ______________________________________________ Nathan C. Nieto, Ph.D., Thesis Advisor ______________________________________________ Tamara Valentine, Ph.D., Director, Honors Program May 2012 3i ABSTRACT This Island Rule is a long-held tenet of biogeography which states that insular populations trend towards increased mean body size in small species and decreased mean body size in large species—a trend that has been inconsistent and even contradictory with reptile taxa. This study examines insular and mainland reptile populations of Aspidoscelis tigris tigris¸ Crotalus oreganus lutosus, Callisaurus draconoides myurus, Sceloporus uniformis, and Sceloporus occidentalis longipes to determine whether the Island Rule conforms with the observed size trends on Anaho Island in Pyramid Lake, Nevada. The selective influences of predation and resource abundance on body size are evaluated by comparing (1) the frequency of caudal autotomy to determine the influence of predation pressure and (2) head shape as a trait affected by the availability of prey resources. Differences in head shape reveal patterns consistent with a shift to smaller prey in A. t. tigris as well as decreased head height for the C. o. lutosus, A. t. tigris, and C. d. myurus. Differences in tail-regeneration frequencies are consistent with an altered pattern of predator-prey interaction for A. t. tigris and S. uniformis. Body size results on Anaho Island contradict the Island Rule, with C. o. lutosus, A. t. tigris, and C. d. myurus males exhibiting smaller body sizes on the island while S. o. longipes and S. uniformis exhibit no size trend, perhaps as the result of a small sample size. Divergence in body size occurs on the island, in a direction that is consistent with the primary literature. This study supports the conclusion that Anaho Island harbors a community of reptiles that is distinct from the mainland in morphology and possibly in ecology and life-history evolution. 4ii ACKNOWLEDGMENTS First and foremost, I would like to thank Drs. Chris R. Feldman and Nathan C. Nieto for their help, their patience, and their unfaltering support. I would also like to thank Tony Bush for providing me with much needed enthusiasm and positivity when I could find none of my own. This research would not have been possible without the contributions of the following individuals and organizations: The California Academy of Sciences, Museum of Vertebrate Zoology, University of Kansas Biodiversity Institute, and San Diego Natural History Museum for providing access to specimens; Xavier Glaudas for generously allowing me to access his data; Drs. Dick Tracy and Chris Gienger for conducting valuable research that served as a strong foundation for this project; Donna Withers of the USFWS, the Nevada Department of Wildlife, and the Pyramid Lake Paiute Indian Tribe for allowing access to study sites and specimens; Peter Murphy for his insightful contributions to statistical analyses; the Honors Undergraduate Research Award (HURA) for providing funding; and Honor’s Program Director Dr. Tamara Valentine who inspires students every day to achieve more than what they think themselves capable of. 5iii TABLE OF CONTENTS ABSTRACT ...................................................................................................................................... i ACKNOWLEDGEMENTS ............................................................................................................ ii TABLE OF CONTENTS ............................................................................................................... iii LIST OF TABLES ......................................................................................................................... iv LIST OF FIGURES ........................................................................................................................ v EPIGRAPH .................................................................................................................................... vi INTRODUCTION .......................................................................................................................... 1 THE ISLAND RULE .................................................................................................................. 1 NATURAL SELECTION: PREDATION PRESSURE ............................................................... 3 NATURAL SELECTION: DIET ................................................................................................ 5 STUDY SITE .............................................................................................................................. 8 SPECIES OF INTEREST .......................................................................................................... 10 OBJECTIVES ........................................................................................................................... 11 MATERIALS AND METHODS .................................................................................................. 14 RESEARCH AREA .................................................................................................................. 14 MEASUREMENT METHODS ................................................................................................ 14 DATA ANALYSIS ................................................................................................................... 16 RESULTS ..................................................................................................................................... 20 BODY SIZE .............................................................................................................................. 20 TAIL-REGENERATION ......................................................................................................... 20 HEAD SIZE .............................................................................................................................. 21 DISCUSSION ............................................................................................................................... 22 BODY SIZE .............................................................................................................................. 22 TAIL-REGENERATION ......................................................................................................... 22 HEAD SIZE .............................................................................................................................. 24 FUTURE DIRECTIONS ........................................................................................................... 25 MANAGEMENT IMPLICATIONS ......................................................................................... 25 WORKS CITED ........................................................................................................................... 27 TABLES AND FIGURES ............................................................................................................ 34 APPENDIX 1 ................................................................................................................................ 45 MUSEUM SPECIMENS .......................................................................................................... 45 6iv LIST OF TABLES Table 1. Results from a Student’s T-test comparing body size between mainland and island individuals of the same sex and species. Average body size (SVL) on Anaho Island relative to Pyramid Lake is significantly different (bolded if P < 0.05) when mean body size of N individuals of differs between locations ......................................................................... 34 Table 2. Overall tail-regeneration frequencies, separated by species for the sexes of either location ....................................................................................................................................................... 35 Table 3. Results from proportion
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