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Proquest Dissertations A STUDY OF THE TIMBER RATTLESNAKE (Crotalus horridus) IN A FRAGMENTED AGRICULTURAL LANDSCAPE A STUDY OF THE TIMBER RATTLESNAKE (Crotalus horridus) IN A FRAGMENTED AGRICULTURAL LANDSCAPE A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biology By Rodney Dale Wittenberg Avila College Bachelor of Science in Biology, 1999 University of Texas at Tyler Master of Science in Biology, 2001 December 2009 University of Arkansas ABSTRACT Few ecological studies have focused on understanding how reptiles use anthropogenically disturbed habitats. Consequently, this research bias has constrained efforts to conserve the ever-increasing number of reptiles threatened by human-induced landscape changes. One such reptile, the Timber Rattlesnake (Crotalus horridus), is a well-studied North American snake species considered to be imperiled by the loss and fragmentation of closed-canopy forest. Despite the threat posed to the Timber Rattlesnake by anthropogenic changes in habitat structure, most research on this species has been carried-out in large tracts of mature forest. Given the important conservation implications of understanding how this species uses anthropogenic habitat, I studied the ecology (spatial, foraging, and thermal) and life history (individual growth rates) of a Timber Rattlesnake population that uses a fragmented agricultural landscape in west- central Missouri. Despite large differences in habitat structure, the spatial data and movement patterns of individuals in this study monitored with radiotelemetry were similar to those previously reported for individuals in closed-canopy forests. However, 9 of the 27 individuals implanted with radiotransmitters (33.3 %) exited the study when they were found dead upon relocation. Sources of mortality were known for 6 of the dead individuals (66.7%), all of which were human-induced. Snakes readily foraged in secondary woodland tracts and corridors, agricultural fields, and habitat edges. Dietary analysis found that snakes fed exclusively on mammals including shrews (Soricidae), mice in the genus Peromyscus, Prairie Voles {Microtus ochrogaster), Cotton Rats (Sigmodon hispidus), Eastern Gray Squirrels (Sciurus caro linens is), and Eastern Cottontails (Sylvilagus floridanus). Although small mammal trapping indicated that fields contained fewer numbers of prey than woodlands and habitat edges, field dwelling Prairie Voles were the most frequently consumed prey item. Data obtained using temperature sensitive radio-transmitters indicated that median snake body temperature (7b) did not differ among woodland (median = 28.0 °C, range = 16.4 - 34.2 °C, n = 63), edge (median = 28.6 °C, range = 16.7 - 33.9 °C, n = 56), or field (median = 28.2 °C, range = 20.2 - 34.4 °C, n = 61) habitats. Additionally, operative temperature models were used to quantify thermal constraints on activity in both a woodland and a field. During each two week interval of the sampling period (02 June to 26 August 2006), the overall percentage of thermally available habitat was high in both woodland (range 84.2 - 99.8 %) and field (range 78.1 - 94.9 %) habitats. However, sharp decreases in habitat availability occurred in both the woodland and field during midday. The lowest hourly percentages of thermally available habitat were recorded at 1400 hours in the field (19.7%) and at 1500 hours in the woodland (36.7%) during the two-week interval from 29 July to 11 August. Body temperatures of gravid females gestating within a man-made quarry (median = 32.5 °C, range = 25.3 - 34.7 °C, n = 113) were significantly warmer and less variable than 7b's of males and non-gravid females using woodlands, edges, and fields (median = 28.3 °C, range = 16.4 - 34.4 °C, n = 180). Finally, mark-recapture data and rattle morphology were used to compare the birth size and early growth rates of timber rattlesnakes in a fragmented habitat of west central Missouri (MO) to those in a closed-canopy forest of northwest Arkansas (AR). Missouri snakes increased in length more rapidly than their AR counterparts through their first eight ecdyses. Furthermore, males and females from MO diverge in size between the fifth and sixth ecdysis event, while growth trajectories of AR males and females remain indistinguishable through eight ecdyses. Despite climatic data suggesting that AR snakes may potentially have an average of 8.5% more time each season to acquire and assimilate prey, MO snakes still exhibited superior growth and early maturation. Results of this study suggest that Timber Rattlesnakes may not require large tracts of closed-canopy forest as long as their thermal and dietary needs are met. Thus, employing habitat management techniques that enhance small mammal densities may be a more effective strategy for conserving Timber Rattlesnake populations than strictly attempting to preserve closed-canopy forest. Future studies should use habitat manipulation as a tool to better understand how forest-dwelling Timber Rattlesnakes respond to changes in habitat structure. This dissertation is approved for Recommendation to the Graduate Council Dissertation Director: Dr. Steven J. Beaupre Dissertation Committee: Dr. Ines Pinto Dr. Kimberly G. Smith Dr. Edward E. Gbur Jr. DISSERTATION DUPLICATION RELEASE I hereby authorize the University of Arkansas Libraries to duplicate this dissertation when needed for research and/or scholarship. Agreed Rodney D. Wittenberg Refused Rodney D. Wittenberg ACKNOWLEDGEMENTS Although few have the opportunity to learn from the best in their respective field, I consider myself "one of the few". I will always be indebted to my friend and advisor, Dr. Steven Beaupre, for generously sharing his time, knowledge, and resources to ensure that I fulfilled a life long dream. I intend to do things in life, personally and professionally, that will make you smile (with pride of course)! As members of my graduate committee, Dr. Edward Gbur Jr., Dr. Ines Pinto, and Dr. Kimberly G. Smith provided guidance and helpful comments on my dissertation. I sincerely appreciate their service. I benefited greatly from interactions with labmates past (Nick Haertl, Dr. Jacques Hill, Dr. Marshal McCue, Carolina Monteiro, Dr. Chad Montgomery, Dr. Melissa Pilgrim, and Dr. Fred Zaidan) and present (Joseph Agugliaro, Lara Douglas, Jason Ortega, Matthew Smith, and James Van Dyke). In one way or another, each of you contributed to my success. For that I thank you. My best friend, Patrick Koontz, provided both assistance and companionship at my study site. Together we flirted with heat stroke, huddled in caves to avoid electrical storms, proved we were immune to insect borne diseases, destroyed several pairs of boots, and had some amazing encounters with the local wildlife. I will always cherish our adventures. I thank my family for their love, support, and encouragement. Extreme examples of their love and support are in order: My father, Robert Wittenberg, allowed me to process my study animals in his spare bedroom. My mother, Peggy Wittenberg, allowed vii me to keep roadkill in the crisper of her refrigerator until it could be preserved or transported elsewhere. Amazing! My grandparents, Virgil and Betty Kempf, have always made me feel special. I thank my Uncle Don and Aunt Diana Wittenberg for always rooting for me. Even before I married my wife, Sara, the Ress family treated me as one of their own. Tom, Roberta, and Michael.. .you have been wonderful. Aside from allowing my study to be conducted on their property, several landowners took a keen interest in me, my project, and the Timber Rattlesnake. For this, I will always be indebted to the Bishop, Debrick, Franklin, Sutton, Thomas, and York families. I am honored to be your friend. I thank the Beckers (Bill and Gwen), the Brewsters (Ambre, Brad, Owen, and Addyson), Adam Crane, Jen Dorr, Matt Dekar, the Ewings (Scott, Linda, Desmond, and Jackson), Andrea Green, Dr. Ron Gutberlet, Ken Hester, Chad Leslie, Jason Luscier, Dr. Glenn Manning, Dr. Nancy McCartney, Alex Muensch, Dr. Douglas James, Dr. Robert Powell, Dana Savorelli, Nora Schubert, the Slays (Mike and Christy), Dr. Sarah Spurrier, and Dr. James Walker. I trust that each of you know why I am saying, "thank you". Finally, this research was approved by the University of Arkansas Institutional Animal Care and Use Committee (protocol # 05001) and the Missouri Department of Conservation (collecting permits # 12005, 12367, 12715, 13101). Partial funding was provided by a University of Arkansas Causey Grant-in-Aid Award, a Harry Steinman Memorial Grant from the St. Louis Herpetological Society, and a grant from the Arkansas Audubon Society Trust. vm DEDICATION I dedicate this dissertation to my wife, Sara (Ress) Wittenberg. Although I have given you an entire page, I am struggling to find the words to fill it. You have enriched my life in every way imaginable, and have done so much to ensure my success. I love you dearly and look forward to the adventures that lie ahead. IX TABLE OF CONTENTS CHAPTER 1. HABITAT FRAGMENTATION AND THE TIMBER RATTLESNAKE (CROTALUS HORRID US): AN INTRODUCTION Abstract 1 Habitat Fragmentation Defined 1 Taxonomic Biases in Habitat Fragmentation Studies 3 The Timber Rattlesnake as a Model for Forest Fragmentation Studies 5 The Timber Rattlesnake in Fragmented Habitats: A Tabular Review 6 Regional Biases 8 Biases Related to Habitat 8 Timber Rattlesnakes in Logged Forests 10 Timber Rattlesnakes in Agricultural Fields 10 Use of Anthropogenic Structures by Timber Rattlesnakes 11
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