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Carabid Beetle Ecology As a Function of Disturbance In CARABID BEETLE ECOLOGY AS A FUNCTION OF DISTURBANCE IN TEMPERATE AND TROPICAL FORESTS BY KATHRYN NICOLE RILEY A Dissertation Submitted to the Graduate Faculty of WAKE FOREST UNIVERSITY GRADUATE SCHOOL OF ARTS AND SCIENCES in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Biology December 2016 Winston-Salem, North Carolina Approved By: Robert A. Browne, Ph.D., Advisor Terry L. Erwin, Ph.D., Chair T. Michael Anderson, Ph.D. William E. Conner, Ph.D. Miles R. Silman, Ph.D. ACKNOWLEDGMENTS First, I want to thank my Advisor Dr. Robert Browne. His guidance, insight and support over the past eight years, not only made this dissertation a success, but profoundly influenced my life. I will be forever grateful for the opportunities in London, Australia and New Zealand which were truly amazing, life enriching experiences. I have been fortunate to work with you all these years and your kindness, patience and advice is sincerely appreciated. I am forever in your debt. Thank you to my committee members, Drs. Miles Silman, T. Michael Anderson, William Conner and Terry Erwin for all of their advice, suggestions and insight during my time as a graduate student, as well as their feedback for this dissertation. I am grateful to Drs. Silman and Anderson from whom I have learned much and who have driven me to be a better ecologist. Thank you, Dr. C for your positive influences over the years and instilling my interest in insects. Terry, to whom I owe so much, I am thankful for the invitations to countless meetings and incorporation into the Carabidologists circle. Thank you for the introduction to Tiputini, one of my favorite places in the world. I cannot thank you enough for the opportunities, insight, knowledge and experiences. Without the support and encouragement of my parents, this degree would not have been possible. Their unconditional support, faith and confidence in my abilities have always encouraged me to pursue my ambitions. To my brother, Nick, for always being there and believing in both me and my goals. Thank you to my family for all the years of stress-filled phone calls, seven hour drives and so much more. To my fiancé, Kevin, thank you for supporting me in all ways possible over the past three and a half years. For ii understanding the many long nights spent in front of the computer. I am forever thankful to have you by my side and look forward to seeing what life has in store for us. I want to mention the professors at Francis Marion University whose influences have led me to this point in my career. Thank you, Travis Knowles for your enthusiasm and influence over the many years since I was an undergraduate in your Organismal Biology class. I appreciate your continued interest and support in my career. To Drs. Tamatha Barbeau and Gregory Pryor, thanks for your support with my undergraduate research project which sparked my interest in field work, diversity and disturbance. I’d like to thank my friends and colleagues, whose support was vital to my success as a graduate student. First thanks to wonderful friends in Maryland, Katie Miller Johnson and Meagan Converse Deal for their continued friendships despite the long distance. Katie, thank you for all long emails, texts and the many, many visits – I am truly lucky to have you as a friend. Thanks to my forever lab-mate, Sarah Maveety, for all the support throughout graduate school as both a lab-mate and friend. Couldn’t have done it without you. To my many friends in the WFU Biology Department (both former and present), thank you for keeping me sane, listening to worries/concerns and, most importantly, all the good times over our many years here. I owe thanks to the undergraduate students who have contributed to these projects over the years. There were many field assistants both in the Amazon and in North Carolina who made the collection of the thousands of carabid beetles possible. I am grateful to the Department of Biology of Wake Forest University for the many years of support, the experiences and the positive environment it provided. iii TABLE OF CONTENTS ACKNOWLEDGEMENTS………………………………………………………….........ii LIST OF TABLES………………………………………………………….……………vii LIST OF FIGURES……………………………………………………………………...xii LIST OF ABBREVIATIONS…………………………………………………………xviii ABSTRACT..…………………………………………………………………………...xix CHAPTER I……………………………………………………………………………….1 General introduction Ecological disturbance and forests……………………………………..…1 Insects and Carabidae…………………………………...………………..2 Objectives……………………..…………………………………………...4 LITERATURE CITED……………………………………………………7 CHAPTER II……………………………………………………………………………..16 Changes in carabid beetle (Coleoptera: Carabidae) assemblages with forest age and sampling year ABSTRACT……………………………………………………………...17 INTRODUCTION……………………………………………………….18 METHODS………………………………………………………………22 RESULTS………………………………………………………………..30 DISCUSSION……………………………………………………………40 ACKNOWLEDGMENTS…..…………………………………………...56 LITERATURE CITED…………………………………………………..57 TABLES…………………………………………………………………79 FIGURES………………………………………………………………...82 iv APPENDIX A……………………………………………………………91 APPENDIX B……………………………………………………………94 APPENDIX C……………………………………………………………98 APPENDIX D…………………………………………………………..107 APPENDIX E…………………………………………………………..109 APPENDIX F…………………………………………………………..111 APPENDIX G…………………………………………………………..117 APPENDIX H…………………………………………………………..120 CHAPTER III…………………………………………………………………………..122 Changes in species traits of carabid beetles (Coleoptera: Carabidae) with forest age ABSTRACT…………………………………………………..………...123 INTRODUCTION…………………………………………………..….124 METHODS……………………………………………………………..128 RESULTS..……………………………………………………………..134 DISCUSSION…………………………………………………..………137 ACKNOWLEDGMENTS…..…………………………………..……...144 LITERATURE CITED…………………………………………………145 TABLES………………………………………………………………..163 FIGURES…………………………………………………………...…..166 APPENDIX A…………………………………………………………..172 APPENDIX B…………………………………………………………..178 APPENDIX C…………………………………………………………..186 v APPENDIX D…………………………………………………………..187 CHAPTER IV…………………………………………………………………………..188 Carabid beetle (Coleoptera: Carabidae) species richness, diversity and community structure in the understory of temporarily flooded and non-flooded Amazonian forests ABSTRACT………………………………………………………….....189 INTRODUCTION……………………………………………………...190 METHODS……………………………………………………………..195 RESULTS………………………………………………………………203 DISCUSSION…………………………………………………………..209 ACKNOWLEDGMENTS…………..………………………………….220 LITERATURE CITED…………………………………………………221 TABLES………………………………………………………………..246 FIGURES……………………………………………………………….248 APPENDIX A…………………………………………………………..257 APPENDIX B…………………………………………………………..259 APPENDIX C…………………………………………………………..265 APPENDIX D…………………………………………………..………270 APPENDIX E………………………………………..…………………272 CHAPTER V…………………………………………………………………………...273 Results from two sampling techniques for carabid beetles (Coleoptera: Carabidae) in temporarily flooded and terra firme rainforest of western Amazonia Published in Studies on Neotropical Fauna and the Environment (2016) vi ABSTRACT………………………………………………………….....274 INTRODUCTION……………………………………………………...275 MATERIALS AND METHODS…...…………………………………..279 RESULTS………………………………………………………………284 DISCUSSION…………………………………………………………..287 CONCLUSIONS AND RECOMMENDATIONS……………………..292 ACKNOWLEDGMENTS……………..……………………………….294 REFERENCES…………………………………………………………295 TABLES………………………………………………………………..310 FIGURES……………………………………………………………….312 APPENDIX 1…………………………………………………………...319 APPENDIX 2…………………………………………………………...322 CHAPTER VI…………………………………………………………………………..324 CONCLUSIONS………………………………...……………………………..324 SCHOLASTIC VITA…………………………………………………………………..326 vii LIST OF TABLES II Table – 1. Measures of abundance, species richness and diversity of Carabidae among the four forest age classes and the summed totals for the dataset…………..………79 II Table – 2. Mean Bray-Curtis dissimilarity values for the forest age classes with all sampling years combined…………………………………………………………...80 II Table – 3. Carabid species with significant associations (p < 0.05) for one of the four forest age classes or groups of forest age classes. Associations were based on the point-biserial group-equalized phi coefficient…………………………….………..81 II Table – 4. Numbers of individuals, species richness and diversity of carabid beetles by sampling year…………………………………………………………………….…82 II Table – B1. Additional site information for the 34 sampling sites including tree circumferences and site elevation…………..……………………………………….94 II Table – B2. Longitude and latitude for the 34 study sites…………………………….97 II Table – C1. Scientific names, species codes and tribes for the 76 Carabidae species collected …………………………….……………………..………………………..98 II Table – C2. Number of individuals of each species collected from the four forest age classes……………………………………………………..……………………….101 II Table – C3. Numbers of individuals for each species collected from the four forest age classes for each sampling year…………………………………………………….103 II Table – D1. Mean Bray-Curtis dissimilarity values by sampling year for community analyses of year-to-year variations among forest age classes……………………..107 II Table – D2. Mean Bray-Curtis dissimilarity by sampling year for community analyses of year-to-year variations among forest age classes ………………………………108 viii II Table – E1. Mean Bray-Curtis dissimilarity values within and among forest age with the first three sampling years combined (extra year of sampling omitted)………..109 II Table – F1. Mean Jaccard dissimilarity values (presence/absence) for the forest age classes with all sampling years combined…………………………………………114 II Table – F2. Mean Jaccard dissimilarity values
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