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Applying Predator-Prey Theory to Evaluate Large Mammal Dynamics

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Applying Predator-Prey Theory to Evaluate Large Mammal Dynamics APPLYING PREDATOR-PREY THEORY TO EVALUATE LARGE MAMMAL DYNAMICS: WOLF PREDATION IN A NEWLY-ESTABLISHED MULTIPLE-PREY SYSTEM by Matthew Smith Becker A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Fish and Wildlife Biology MONTANA STATE UNIVERSITY Bozeman, Montana December 2008 ©COPYRIGHT by Matthew Smith Becker 2008 All Rights Reserved ii APPROVAL of a dissertation submitted by Matthew Smith Becker This dissertation has been read by each member of the dissertation committee and has been found to be satisfactory regarding content, English usage, format, citation, bibliographic style, and consistency, and is ready for submission to the Division of Graduate Education. Dr. Robert A. Garrott, Committee Chair Approved for the Department of Ecology Dr. David Roberts, Department Head Approved for the Division of Graduate Education Dr. Carl. A. Fox, Vice Provost iii STATEMENT OF PERMISSION TO USE In presenting this dissertation in partial fulfillment of the requirements for a doctoral degree at Montana State University, I agree that the Library shall make it available to borrowers under rules of the Library. I further agree that copying of this dissertation is allowable only for scholarly purposes, consistent with “fair use” as prescribed in the U.S. Copyright Law. Requests for extensive copying or reproduction of this dissertation should be referred to ProQuest Information and Learning, 300 North Zeeb Road, Ann Arbor, Michigan 48106, to whom I have granted “the exclusive right to reproduce and distribute my dissertation in and from microform along with the non- exclusive right to reproduce and distribute my abstract in any format in whole or in part.” Matthew Smith Becker December 2008 iv ACKNOWLEDGMENTS A one-page limit sadly precludes adequate recognition of many essential contributions to this project; more appropriate acknowledgements can be found in the published work. Major research funding was provided by a National Science Foundation Graduate Research Fellowship a GK-12 Fellowship and NSF grants (DEB-9806266, DEB-0074444, DEB-0413570, DEB-0716188) to Robert A. Garrott, Lee Eberhardt, and P. J. White. The National Park Service made this research possible with their assistance in all facets of the work. Thanks to my advisor R. Garrott for his mentoring and generosity in providing me this opportunity and to committee members J. Borkowski, S. Creel, S. Kalinowski, and P. J. White for providing useful input and advice during every phase of this research. This work benefited immensely from reviews from Bruce Dale. Thanks to all field crews from 1996-97 to 2006-07, with special gratitude to R. J., E. Bergman, C. Gower and S. Dunkley for essential contributions and continuity as students and crew leaders. To S. Dunkley for keeping things running under extremely difficult circumstances, and to the memory of her mother, Andra, who passed away during this time. Special thanks to all my great co-workers in the field: J. Bruggeman, S. Dunkley, C. Gower, V. Green, C. Kenyon, K. Pietrzak, E. Robertson, T. Shafer, D. Thompson, and V. Yovovich and to my fellow students for their advice, humor, and support. Personal support was provided by my loving family: my parents, Michael and Stephanie, my sister Clare, late dog Maud, step-dogs Bijou and Paco, and my wonderful friends. Special thanks also go to my dear dad for nurturing and inspiring my childhood love of wild animals and wild places and for encouraging me to pursue a difficult but fulfilling path. v TABLE OF CONTENTS 1. INTRODUCTION TO DISSERTATION ...................................................................... 1 Introduction..................................................................................................................... 1 Study Area ......................................................................................................................4 Literature Cited ............................................................................................................... 8 2. WOLF PREY SELECTION IN AN ELK-BISON SYSTEM: CHOICE OR CIRCUMSTANCE?..................................................................................................... 12 Introduction................................................................................................................... 12 Methods......................................................................................................................... 18 Detecting and Identifying Wolf Kills ....................................................................... 18 Factors Influencing Wolf Prey Selection.................................................................. 21 Prey Switching.......................................................................................................... 30 Results........................................................................................................................... 33 Factors Influencing Wolf Prey Selection.................................................................. 42 Wolf Prey Switching with Murdoch’s Equation....................................................... 44 Discussion..................................................................................................................... 47 Literature Cited ............................................................................................................. 61 3. WOLF KILL RATES: PREDICTABLY VARIABLE?............................................... 73 Introduction................................................................................................................... 73 Methods......................................................................................................................... 77 Wolf Tracking and Kill Detection ............................................................................ 77 Kill Rate Estimation.................................................................................................. 79 Evaluating Kill Rate Variation.................................................................................. 80 Evaluating the Functional Response for Elk............................................................. 83 Results........................................................................................................................... 86 Variations in Elk Kill Rate........................................................................................ 91 Variations in Bison and Total Kill Rates .................................................................. 97 Discussion................................................................................................................... 101 Literature Cited ........................................................................................................... 115 4. SYNTHESIS AND DIRECTIONS FOR FUTURE RESEARCH ............................. 125 Synthesis ..................................................................................................................... 125 Management and Conservation Implications and Directions for Future Research ........................................................................ 128 Literature Cited ........................................................................................................... 132 vi TABLE OF CONTENTS-CONTINUED APPENDICES ................................................................................................................ 134 APPENDIX A: Multinomial Model List ................................................................... 135 APPENDIX B: A Priori Model Lists for Evaluating Kill Rate Variation ................. 140 APPENDIX C: Description of Multiple Species Functional Response Models........ 145 vii LIST OF TABLES Table Page 2.1. Numbers of bison and elk carcasses killed by wolves and detected in the Madison headwaters area of Yellowstone National Park during 1996-97 through 2006-07....................................................................................36 2.2. Selection indices (Chesson 1978) for four prey types from resident wolf pack kills during winters 1998-99 through 2006-07...................................39 2.3. A priori model structure and results from top models within and among suites for multinomial logit analyses of winter prey selection by resident wolf packs in a bison-elk system in the Madison headwaters area of Yellowstone National Park during 1998-99 through 2006-07. ...........................43 2.4. Models identified through AIC model comparison techniques for prey selection by resident wolf packs on bison and elk in the Madison headwaters area of Yellowstone National Park during 1998-99 through 2006-07....................................................................................46 2.5. Coefficient values and lower and upper 95% confidence intervals from analyses of prey switching by wolves in a bison-elk system in the Madison headwaters area of Yellowstone National Park during 1998-99 through 2006-07....................................................................................47 3.1. A priori model structure and results from top models for multiple linear regression analyses of elk kill rates by resident wolf packs in the Madison headwaters area of Yellowstone National Park during 1998-99 through 2006-07..................................................................................................93 3.2. A priori model structure and results from top models for multiple linear
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