WOLVES, COYOTES, ELK, and MULE DEER: PREDATOR-PREY BEHAVIORAL INTERACTIONS in SOUTHWESTERN MONTANA by Todd C. Atwood a Dissertat
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WOLVES, COYOTES, ELK, AND MULE DEER: PREDATOR-PREY BEHAVIORAL INTERACTIONS IN SOUTHWESTERN MONTANA by Todd C. Atwood A dissertation submitted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY in Wildlife Biology Approved: Dr. Eric Gese Dr. Kyran Kunkel Major Professor Committee Member Dr. Fred Provenza Dr. Robert Schmidt Committee Member Committee Member umham Committee Member iuate Studies UTAH STATE UNIVERSITY Logan, Utah 2006 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. UMI Number: 3252259 INFORMATION TO USERS The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleed-through, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. ® UMI UMI Microform 3252259 Copyright 2007 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Copyright © Todd Atwood 2006 All Rights Reserved Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Wolves, Coyotes, Elk, and Mule Deer: Behavioral Interactions in Southwest Montana by Todd C. Atwood, Doctor of Philosophy Utah State University, 2006 Major Professor: Dr. Eric M. Gese Department: Wildland Resources Predator recolonization can have profound effects on prey communities, particularly when a predator recolonizes an already complex predator-prey community. In the following chapters, I investigated direct and indirect effects of wolf ( Canis lupus) recolonization on prey and competitor behavior. In Chapter II, I examined comparative patterns of cougar ( Puma concolor) and wolf predation on elk (Cervus elaphus), mule deer ( Odocoileus hemionus), and white-tailed deer ( Odocoileus virginianus). I found that increases in the proportion of elk killed by cougars were mediated by wolves inducing elk to shift to more structurally complex refugia habitat to mitigate predation risk from wolves which, in turn, made elk more vulnerable to predation by cougars. In Chapter III, I examined the interaction between landscape attributes and predator and prey resource selection in influencing the spatial distribution of predation risk. Using the functional response theoretic to model total predation risk for elk and mule deer, I found that landscape covariates could exert a strong influence in mediating the conditional probability of death following a predator encounter. In Chapters IV and V, I investigated the effects of wolf recolonization on coyote ( Canis latrans) space use and scavenging Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. behavior. I found that coyotes routinely exploit space used by wolves putatively in the hopes of encountering scavenge subsidies in the form of wolf-killed prey. Upon encountering wolf-killed prey, coyotes rely on a gradient of risk-sensitive behaviors, ranging from increased vigilance to aggressively confronting wolves, to guide scavenging behavior. Coyote social rank strongly influences risk sensitivity as socially dominant coyotes spend more time vigilant at carcasses, but also are exposed to greater risk as they tend to feed at carcasses in the earliest stages of consumption by wolves. As a result, there is greater potential for agonistic interactions between socially dominant coyotes and wolves. (165 pages) Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. V DEDICATION To my wife Laura, we’ve done it! For the wonderful canine companions that sparked and maintain my interest in canid behavior- thank you Sid, Joker, Beverly, Orson, Carmilla, Ike, Midge, Monkey, and Blue. I’ve had a joyful time studying your canid cousins. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGMENTS I would like to thank Dr. Eric Gese and my committee members, Dr. Kyran Kunkel, Dr. Fred Provenza, Dr. Robert Schmidt, and Dr. Mike Wolfe for support and assistance throughout the dissertation process. I am grateful for the financial support provided by the National Wildlife Research Center, Boone and Crockett Club, Utah State University Presidential Fellowship Program, and Pope and Young Club. Turner Enterprises, Inc., and the Turner Endangered Species Fund assisted with logistical support. B. Beehler, J. Brown, A. Corcoran, A. Nelson, and K. Nittinger provided valuable assistance in the field. I thank V. Asher and M. Phillips of the Turner Endangered Species Fund, M. Kossler of Turner Enterprises, Inc., R. Amaud and J. Doran of Montana Outfitters, and D. Pac of Montana Fish, Wildlife, and Parks for their advice and help with various aspects of this research. I thank T. Turner and B. Turner for providing access to the NMSA. I owe a debt of gratitude to Mike Ebinger, Julie Young, Kim Berger, and Pat Terletzky for numerous discussions that informed various aspects of this research. Finally, this work would not have been possible without the emotional and financial support of my wife, Laura, and the encouragement of Peggy and Bernard Speranza and David and Lisa Lautz. Research and handling protocols were reviewed and approved by Institutional Animal Care and Use Committees at the National Wildlife Research Center and Utah State University. Todd Atwood Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CONTENTS Page ABSTRACT...................................................................................................................... iii DEDICATION.................................................................................................................... v ACKNOWLEDGMENTS................................................................................................. vi LIST OF TABLES..............................................................................................................ix LIST OF FIGURES............................................................................................................xi CHAPTER I. INTRODUCTION..............................................................................................1 References.........................................................................................................5 II. COMPARATIVE PATTERNS OF PREDATION BY COUGARS AND RECOLONIZING WOLVES IN MONTANA’S MADISON RANGE..........8 Introduction ......................................................................................................8 Methods ...........................................................................................................11 Results.............................................................................................................17 Discussion ........................................................................................................21 References.......................................................................................................26 III. SPATIAL PARTITIONING OF TOTAL PREDATION RISK IN A MULTIPLE-PREDATOR MULTIPLE-PREY SYSTEM.............................37 Introduction .....................................................................................................38 Methods ...........................................................................................................42 Results.............................................................................................................47 Discussion .......................................................................................................51 References.......................................................................................................58 IV. RESOURCE SELECTION AND SOCIAL BEHAVIOUR MODULATES THE PARTITIONING OF HOSTILE SPACE BY SYMPATRIC CANIDS..........................................................................................................72 Introduction .....................................................................................................73 Methods ...........................................................................................................78 Results.............................................................................................................83 Discussion .......................................................................................................86 References.......................................................................................................91 V. SCAVENGING ECOLOGY AND RECOLONIZING WOLVES: FEAR Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. AND LOATHING IN MONTANA’S MADISON RANGE?...................... 108 Introduction .................................................................................................. 109 Methods .......................................................................................................