WHITE-TAILED DEER, COYOTES, AND THE ECOLOGY OF FEAR IN A LONGLEAF PINE SAVANNA By MICHAEL JOHN CHERRY (Under the Direction of Robert J. Warren) Abstract Predators can exert powerful influence on their prey, independent of direct killing, by inducing antipredator responses. Coyotes (Canis latrans) have recently achieved abundances capable of influencing white-tailed deer (Odocoileus virginianus) population demography in the southeastern USA, but the effects of antipredator responses have not been reported. I conducted a multifaceted investigation of coyote and white-tailed deer interactions, using population monitoring data, harvest data, and results from controlled experimentation with predator exclosures. This work provided evidence that coyotes can influence white-tailed deer space use and vigilance while foraging, and documented a negative relationship between coyote abundance and body mass of adult female deer during an 11-year period. I compared multiple measures of reproductive success during a 7-year period that encompassed high and low coyote-deer ratios to elucidate the relative contributions of direct predation and predation risk effects to an observed increase on recruitment as measured by fawn-adult female ratios. Fawn survival rates were similar between periods, but the proportion of females with evidence of ovulation increased during the period of low coyote-deer ratios. Increases in ovulation were similar to increases in the proportion of females with evidence of lactation and fawn-adult female ratios. While direct killing by predators greatly influenced survival of fawns during both periods, changes in recruitment resulted from variations in fecundity. I tested hypotheses predicting the consequences of 10 years of predator exclusion on oak (Quercus sp.) recruitment and the density of selected deer forage species. Oaks are an important component of the longleaf pine savannas, and factors influencing their recruitment are of significant importance to the restoration and management of the longleaf pine (Pinus palustrus) ecosystem. Predator exclusion increased oak recruitment and decreased the density of selected deer forage species in a frequently burned longleaf pine savanna. Thus, coyote predation risk can influence white-tailed deer herbivory and thereby potentially affect composition of groundcover and hardwood understory in longleaf pine ecosystems. This study demonstrates that coyotes can have strong predation risk effects on white-tailed deer populations, and ignoring these effects may result in dramatic underestimation of impacts of expanding coyote populations on ecosystems. Index words: Body mass, coyote, Canis latrans, fecundity, herbivory, longleaf pine, non- consumptive effects, Odocoileus virginianus predation risk effects, recruitment, trophic cascade, vigilance, white-tailed deer WHITE-TAILED DEER, COYOTES, AND THE ECOLOGY OF FEAR IN A LONGLEAF PINE SAVANNA By MICHAEL JOHN CHERRY B.S.F.R. University of Georgia 2010 A Dissertation Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY Athens, Georgia 2014 © 2014 Michael John Cherry All Rights Reserved WHITE-TAILED DEER, COYOTES, AND THE ECOLOGY OF FEAR IN A LONGLEAF PINE SAVANNA by MICHAEL JOHN CHERRY Major Professor: Robert J. Warren Committee: L. Mike Conner Nathan P. Nibbelink Lindsay R. Boring Electronic Version Approved: Julie Coffield Interim Dean of the Graduate School The University of Georgia August 2014 DEDICATION I dedicate this dissertation to my wife, Taulbee, and son, John Russell, for providing the right amount of motivation and distraction throughout the process. iv ACKNOWLEDGEMENTS Funding for this project was provided by the Georgia Wildlife Resources Division (GA-WRD) through the Wildlife Restoration Program. This program is funded through the Pittman– Robertson Federal Aid in Wildlife Restoration Act which places an excise tax on sporting arms and ammunition paid by hunters and recreational shooters. I am deeply indebted to the sportsman and women that enjoy the wildlife resources and recreational activities in the state of Georgia. I thank my collaborators at the GA-WRD, Charlie Killmaster and John Bowers. Charlie’s support for applied research and his desire to integrate the latest science into decision making processes provided motivation and inspiration during this research. I thank John Bowers for providing a view into complexities of managing wildlife resources while considering the position of multiple stake holders. This view has helped shaped my perspective on the role of science in policy and management. I thank the GA-WRD for the having the foresight to be the first southeastern state to investigate the non-consumptive effects of coyotes, demonstrating the GA-WRD’s dedication to being at the forefront of applied research and management. I was fortunate to be co-advised by Dr. Bob Warren and Dr. Mike Conner whose collaboration provided a model of effective and synergistic professional partnership. Bob Warren is my professional model and mentor. I appreciate his constant encouragement and positive approach to work and life. Mike Conner is largely responsible for my development as a research scientist. I have distracted him for countless hours discussing differences between “what we know” and “what we can show” about our study system. Much of my development v occurred in the offices of Bob Warren and Mike Conner. I thank my advisory committee members, Dr. Nate Nibbelink and Dr. Lindsay Boring for encouraging me to ask questions at multiple scales and ecological levels of organization. I am fortunate to have had access such insightful scientific advisement, and I have gained tremendously through discussions with Nate Nibbelink and Lindsay Boring. Many other scientists have been influential in my development. Dr. Steven Castleberry has had a tremendous influence at many stages of my professional development; he has been my instructor, professional advisor, instructional mentor, and friend. I thank him for his friendship and contribution to my professional development. I thank Dr. Karl V. Miller for sharing his vast expertise in white-tailed deer ecology and management, and for providing me countless opportunities to disseminate my research to land managers. I thank Dr. Mike Chamberlain for his input into this project and for sharing his wealth of knowledge on coyote ecology. I thank Dr. Mike Mengak for his encouragement and facilitation of my involvement with the Wildlife Society and particularly the Wildlife Damage Management Working Group. I thank Dr. John Carroll for our many discussions and his candid prospective on science, academia, and longleaf pine management. I thank Dr. Kay Kirkman for providing her expertise in plant ecology, collaboration, and honest assessment of my hypotheses. I thank Dr. Lora Smith for her collaboration and for sharing her knowledge of the ecology of longleaf pine ecosystem. I thank Dr. Liana Zanette for providing a review of part of this dissertation and for encouraging me to consider the broader ecological relevance of my results. Dr. Joel Brown’s theoretical framework for the ecology of fear was inspirational to this dissertation, and I am truly gratefully that the Jones Center hosted Joel and provided me the opportunity to discuss my research with him standing in a longleaf pine woodland. vi I consider myself very fortunate to have had the opportunity to conduct my dissertation research on Joseph W. Jones Ecological Research Center at Ichauway. I am deeply indebted to the Jones Center and the Robert W. Woodruff Foundation. Many people on the Jones Center contributed to this research. I thank Jean Brock for her help with spatial analysis, encouragement, and friendship. Brandon Rutledge is a valuable collaborator who was fundamental in this research. Michael Simmons facilitated data handling and analysis. His contributions were invaluable and saved the project hundreds of hours of labor. I thank Bobby Bass and Jerry Wade for sharing their knowledge of Ichauway and for help capturing animals. I thank all the folks that helped capture fawns, who are too numerous to list here. I thank Chaz Oliver, John-Michael McCormick, Austin Baggarly, Rob Dukes, and Joe Warden for data collection and maintaining a positive attitude during adverse conditions. I thank the students who conducted their Senior Thesis research in conjunction with this project including, Kim Alles, Keri Morgan, Cody Seagraves, Clay Stroud, and Kelsey Turner. I thank Melinda Nelson for her work on this project and all the members of UGA Deer Lab and the Jones Center Wildlife Ecology Lab for support. vii TABLE OF CONTENTS Page ACKNOWLEDGEMENT ...............................................................................................................v LIST OF TABLES ......................................................................................................................... xi LIST OF FIGURES ..................................................................................................................... xiv CHAPTER 1 INTRODUCTION, LITERATURE REVIEW, STUDY AREA, DISSERTATION FORMAT .................................................................................................................1 INTRODUCTION ...................................................................................................1 LITERATURE REVIEW ........................................................................................3 STUDY AREA