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Etd-06032014-173652.Pdf Automated Template B: Created by James Nail 2011V2.1 Expression of facultative den behavior in Ursidae By Nicholas Lee Fowler A Thesis Submitted to the Faculty of Mississippi State University in Partial Fulfillment of the Requirements for the Degree of Master of Science in Wildlife, Fisheries and Aquaculture in the Department of Wildlife, Fisheries and Aquaculture Mississippi State, Mississippi August 2014 Copyright by Nicholas Lee Fowler 2014 Expression of facultative den behavior in Ursidae By Nicholas Lee Fowler Approved: ____________________________________ Jerrold L. Belant Major Professor ____________________________________ Bruce D. Leopold Committee Member ____________________________________ Guiming Wang Committee Member ____________________________________ Eric D. Dibble Graduate Coordinator ____________________________________ George M. Hopper Dean College of Forest Resources Name: Nicholas Lee Fowler Date of Degree: August 15, 2014 Institution: Mississippi State University Major Field: Wildlife, Fisheries and Aquaculture Major Professor: Jerrold L. Belant Title of Study: Expression of facultative den behavior in Ursidae Pages in Study: 60 Candidate for Degree of Master of Science Many ecological and evolutionary processes have been described underlying natural selection but supporting evidence in wild settings is often lacking and our understanding remains incomplete. Using a multi-scale approach, I investigated expression of facultative den behavior (denning) among and within ursid species. Among species, lack of denning behavior was supported by relaxation of selection pressures of seasonal food availability due to realization of two sources of ecological opportunity; colonization of new habitat and adaptation of key innovations. Of denning species, I found evidence of adaptive ecological plasticity in den chronology among and within populations consistent with relaxed selective pressures influencing seasonal food availability and energetic budgets. I supported these finding using indices of fitness and established ecological theory. Further knowledge of organismal response to selection pressures has practical applications in predicting responses to novel and fluctuating pressures and aids in our understanding of the complexities of ecology and evolution. DEDICATION For their endless support and encouragement, I dedicate this thesis to my family. ii ACKNOWLEDGEMENTS I appreciate the support of many people for their contributions to this thesis and the field component of my master’s program. I thank Dr. Jack Jones and Dr. Josh Millspaugh at Mizzou for encouraging me to pursue graduate studies. I thank Jerry for bringing me on board and his unwavering patience, support, and encouragement every step of the way. I thank my committee members, Dr. Bruce Leopold and Dr. Guiming Wang, for sharing their knowledge of ecological research with me. I thank Dr. Sam Riffell and Dr. James Martin for assistance in statistical analysis. I thank the members of the Carnivore Ecology Lab and other departmental graduate students for providing a springboard for ideas and the occasional distraction. Additionally, I appreciate the work of all authors, institutions, and agencies whose published data was used in this project and Dr. Larry Van Daele and Vic Barnes for unpublished data. I thank Danny Gammons for my first experiences capturing bears, and Brad Young for expanding and refining these skills. I thank the Mississippi Department of Wildlife, Fisheries, and Parks and other state and federal agencies and private organizations for their roles in the Mississippi Black Bear Project. I thank the departmental administrative staff for keeping everything running as smooth as possible. Last but not least, I thank the Delta landowners and all interested folks for making my field work such a fulfilling and enjoyable experience. iii TABLE OF CONTENTS DEDICATION .................................................................................................................... ii ACKNOWLEDGEMENTS ............................................................................................... iii LIST OF TABLES ............................................................................................................. vi LIST OF FIGURES .......................................................................................................... vii CHAPTER I. EXPRESSION OF FACULTATIVE DEN BEHAVIOR SUPPORTS ECOLOGICAL OPPORTUNITY IN URSIDS .....................................1 Introduction ........................................................................................................1 Colonization of new habitat ...............................................................................3 Seasonal and annual diet ..............................................................................4 Climate seasonality ......................................................................................6 Key innovations .................................................................................................7 Trait adaptations...........................................................................................8 Energetics of denning ..................................................................................9 Conclusion .......................................................................................................12 References ........................................................................................................17 II. RELAXATION OF SELECTIVE PRESSURES AND ECOLOGICAL PLASTICITY IN URSID DEN BEHAVIOR......................................24 Introduction ......................................................................................................24 Methods............................................................................................................26 Inter-population plasticity ..........................................................................28 Intra-population plasticity ..........................................................................30 Fitness related effects .................................................................................30 Results ..............................................................................................................32 Inter-population plasticity ..........................................................................32 Intra-population plasticity ..........................................................................33 Fitness related effects .................................................................................34 Discussion ........................................................................................................34 Inter-population plasticity ..........................................................................35 Intra-population plasticity ..........................................................................38 Conclusion .......................................................................................................39 iv References........................................................................................................48 III. SUMMARY AND CONCLUSION ................................................................58 References........................................................................................................60 v LIST OF TABLES 2.1 Global models and model term descriptions used in analyses of factors influencing den chronology and variability in den chronology of American black bear and brown bear in North America........................40 2.2 Best supported model (∆AICc ≤ 2) results of factors influencing den chronology and variability of den chronology of American black bear and brown bear in North America. .................................................41 2.3 Model averaged parameter estimates of factors influencing den chronology and variability in den chronology for American black bear and brown bear in North America. .......................................42 vi LIST OF FIGURES 1.1 Generalized global distributions of bear species. Continental boundaries and distribution of Andean bear and giant panda smoothed for visualization (data from IUCN red list)..................................................14 1.2 Global comparison of annual mean temperature seasonality, precipitation seasonality, and temperature of annual quarter (left panels). Statistical characteristics of these climate indices across distributions of exant ursids (right panels).............................................15 1.3 Relationship between active and denning minimum energetic requirements of ursid spp. over mean duration of denning of non-parous American black bears (137 days; Fowler et al. 2014).........16 2.1 Locations of American black bear (circles) and brown bear (squares) studies reporting den chronology used in analyses. ...............................43 2.2 Mean den chronology and associated variability of American black bear (circles) and brown bear (squares) against latitude in North America ..................................................................................................44 2.3 Mean effect size (+ 95% CI) of experimental groups: parturient females (PF), non-parturient females (NPF), and sub adults (SA) den entrance date and duration of denning relative to control group, adult males (AM) of American black bear (squares), brown bear (circles), and species combined (triangles) in North America. Numbers represent sample sizes.
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