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Implications for Pronghorn Persistence in Grand Teton CONSERVATION IMPLICATIONS OF FOOD WEBS INVOLVING WOLVES, COYOTES, AND PRONGHORN by Kim Murray Berger A dissertation submitted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY in Wildlife Biology UTAH STATE UNIVERSITY Logan, Utah 2007 ii Copyright © Kim Murray Berger 2007 All Rights Reserved iii ABSTRACT Conservation Implications of Food Webs Involving Wolves, Coyotes, and Pronghorn by Kim Murray Berger, Doctor of Philosophy Utah State University, 2007 Major Professor: Eric M. Gese Department: Wildland Resources Coyote (Canis latrans) predation is an important factor in the mortality of neonatal pronghorn (Antilocapra americana). It has been suggested that the extirpation of gray wolves (Canis lupus) contributed to an overall increase in coyote densities and a concomitant rise in predation rates on pronghorn fawns, a process known as trophic cascades. To test this hypothesis we contrasted cause-specific mortality and survival rates of fawns captured at wolf-free and wolf-abundant sites; used demographic modeling to assess the impact of wolves on pronghorn population dynamics; and evaluated coyote survival rates, causes-specific mortality, and spatial segregation with wolves, to identify mechanisms by which wolves limit coyote densities. Fawn survival rates were 400% higher, and coyote densities 33% lower, at sites used by wolves. Wolves killed 56% of transient coyotes, and dispersal rates of transients were 117% higher at the wolf-abundant site; thus, differential effects on solitary coyotes may be an important mechanism by which wolves reduce coyote densities. Our results support the hypothesis that the iv extirpation of wolves contributes to high rates of coyote predation on pronghorn fawns, and add to a growing body of evidence demonstrating the importance of top-down forces in structuring the dynamics of consumer-resource interactions. (169 pages) v ACKNOWLEDGMENTS Funding for this research was provided by Grand Teton National Park and the Biological Resources Division of the U.S. Geological Survey under Cooperative Agreement No. 01CRAG0031, with additional support from the Wildlife Conservation Society, National Wildlife Research Center at Utah State University, NatureFlight Foundation, and Earth Friends Foundation. I thank Noah Weber, Renee Wulff, Mike Nordell, Jana Martinez, Sonja Berger, and the pronghorn capture team volunteers for field assistance, and Jim Pehringer (Wildlife Services) for assistance with coyote captures. Grand Teton National Park, the Wyoming Game & Fish Department, the U.S. Fish & Wildlife Service, and the U.S. Forest Service granted necessary permits. Mike Jimenez, Sarah Dewey, and Lisa Robertson provided wolf locations. I am indebted to Mary Conner for her infinite patience and assistance with issues related to Program MARK, simulation modeling, and statistical analysis, and Gary White for indulging all my “true or false” questions about random effects models and variance components. John Fieberg graciously provided code for the analysis of home-range overlap. I thank Eric Gese for his professionalism throughout the graduate school process, for allowing me to pursue my intellectual interests while still providing guidance when I needed it, and for somehow finding time to assist with field work every year. John Bissonette, Mary Conner, Dan Rosenberg, and Phaedra Budy provided thoughtful comments on my manuscripts. Lastly, I thank Joel for being a source of inspiration, for his patience during my long absences from home, and for being my biggest fan. Kim Murray Berger vi PREFACE Chapter 2, Does Interference Competition with Wolves Limit the Distribution and Abundance of Coyotes?, will be submitted to the Journal of Animal Ecology. The sole co-author will be Eric M. Gese. Chapter 3, Evidence of a Species-Level Trophic Cascade in a Terrestrial Food Web Involving Wolves, Coyotes, and Pronghorn, will be submitted to Ecology. Coauthors will be Eric M. Gese and Joel Berger. Chapter 4, Recolonizing Wolves and Mesopredator Suppression of Coyotes: Impacts on Pronghorn Population Dynamics, will be submitted to Ecological Applications. The sole co-author will be Mary M. Conner. vii CONTENTS Page ABSTRACT....................................................................................................................... iii ACKNOWLEDGMENTS ...................................................................................................v PREFACE.......................................................................................................................... vi LIST OF TABLES............................................................................................................. ix LIST OF FIGURES ........................................................................................................... xi CHAPTER 1. INTRODUCTION .......................................................................................1 2. DOES INTERFERENCE COMPETITION WITH WOLVES LIMIT THE DISTRIBUTION AND ABUNDANCE OF COYOTES?..................6 3. EVIDENCE OF A SPECIES-LEVEL TROPHIC CASCADE IN A TERRESTRIAL FOOD WEB INVOLVING WOLVES, COYOTES, AND PRONGHORN.................................................................................45 4. RECOLONIZING WOLVES AND MESOPREDATOR SUPPRESSION OF COYOTES: IMPACTS ON PRONGHORN POPULATION DYNAMICS ....................................................................84 5. CONCLUSIONS......................................................................................130 APPENDICES .................................................................................................................137 Appendix A. Coyote Densities Predicted by Scat Deposition Index ................138 Appendix B. Description of Known Fate Models Examined in the Analysis of Neonatal Pronghorn Survival ..................................................140 Appendix C. Model-Selection Results Comparing the Performance of Models by Site and Treatment and Resulting Estimates of Neonatal Survival. ...................................................................142 Appendix D. A Technique to Estimate the Number of Reproductive Females in the Population and Neonatal Survival Rates from Count Data ..........................................................................145 Appendix E. Effect of Percentage Yearlings in the Initial Population Vector on the Results of Demographic Modeling ...................................147 viii Appendix F. Effect of Correlation in Overwinter Survival Rates on the Results of Demographic Modeling ..............................................150 Appendix G Permission-to-Use Letters............................................................152 CURRICULUM VITAE..................................................................................................154 ix LIST OF TABLES Table Page 2-1 Model selection results for survival of coyotes at two study sites in Grand Teton National Park, Wyoming, 2001-2004...............................................37 2-2 Seasonal and annual estimates of coyote survival at two study sites in Grand Teton National Park, Wyoming, 2002-2004..........................................................38 2-3 Population densities of coyotes and wolves at three study areas in the Greater Yellowstone Ecosystem, 1991-2005.........................................................39 2-4 Overlap between annual coyote and seasonal wolf home ranges and core areas as measured by percentage of area and utilization distribution overlap index (UDOI) in Grand Teton National Park, Wyoming, 2002-2004.............................40 3-1 Descriptions of variables included in the analysis of pronghorn fawn survival....75 3-2 Model selection results for survival of pronghorn fawns during the first 60 days of life at three study sites in northwestern Wyoming, 2002-2004.................76 3-3 Model-averaged estimates of pronghorn fawn survival during the first 60 days of life at three study sites in northwestern Wyoming, 2002-2004.........................77 4-1 Comparison of observed neonatal survival rates of radio-collared fawns and neonatal survival rates estimated from young:female ratios during fall classification counts.............................................................................................117 4-2 Stage classes, ages, and vital rates used to construct the demographic models...118 4-3 Survival rates for adults and juveniles used to estimate correlation in overwinter survival ..............................................................................................119 4-4 Average population projection matrices for pronghorn based on differences in survival rates of neonates at wolf-free and wolf-abundant sites in Grand Teton National Park, Wyoming ...........................................................................120 4-5 Sample sizes (n) and percentages of radio-collared pronghorn fawns dying by various causes during the first 60 days of life in Grand Teton National Park, Wyoming. ...................................................................................................121 4-6 Model selection results from analysis of neonatal pronghorn survival at wolf- free and wolf-abundant sites in Grand Teton National Park, Wyoming..............122 x 4-7 Comparison of estimated survival rates for male and female fawns based on parameter estimates from the top-ranked model..................................................123 A-1 Results of regression analysis to estimate coyote densities as a
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