ECOLOGY OF GROUND-DWELLING CARABID BEETLES (COLEOPTERA: CARABIDAE) IN THE PERUVIAN ANDES BY SARAH A. MAVEETY 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 2013 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 would like to thank to my advisor, Dr. Robert Browne, who has played a pivotal role in my time at Wake Forest University, encouraging and supporting my research endeavors since my sophomore year as a biology undergraduate. I would also like to thank Biology Department committee members, Dr. Miles Silman, Dr. Bill Conner, Dr. Michael Anderson. The dissertation research would not have taken place in Peru had it not been for Dr. Silman’s long standing research presence there, and I am grateful for his connection. Dr. Conner was my first professor in biology ten years ago, and he has given invaluable insight as the first reader of the dissertation. Dr. Anderson provided instrumental ecological perspective in both my academic career and dissertation. A special thanks to my outside committee member, Dr. Terry Erwin, Curator of Coleoptera at the Smithsonian National Museum of Natural History, and Neotropical carabid beetle expert. He has dedicated his time and expertise so that my specimens were identified correctly and I made all the appropriate connections to colleagues in both Peru and the U.S. The field portion of this project was supported by a Fulbright Grant to conduct research in Peru, and I thank the onsite staff of the Comisión de Fulbright in Lima, Peru. I am also grateful to colleagues at the Museo de Historia Natural Universidad Nacional Mayor de San Marcos, Dr. Gerardo Lamas, Juan Grados, and Luis Figueroa, for help in obtaining permits for beetle collection and exportation. For assistance with field logistics I’d like to thank: Asocación para la Conservación de la Cuenca Amazonica (ACCA); Daniel Blanco at PeruVerde; ProNaturaleza; students and professors at Universidad Nacional San Antonio del Abad Cusco and the Museo de Historia Natural, especially my ii field assistants Jessica Ttito Quispe and Carla Chaparro Zamalloa; and Hubert Jaquehua Callo, No lo hubiera logrado sin ti. Finally, I want to thank former Browne lab graduate students Carmen Chavez and Doug Bruce for sharing Appalachian ground beetle data, and to all the undergraduates who helped in data collection and helped me to meticulously measure individual body lengths. Thank you to my all friends at Wake Forest University, but especially to Rachel Hillyer for her friendship and for help studying for preliminary exams, and to Katie Riley, for going through the ups and downs of graduate school as both a best friend and colleague. I thank my best friends from home, Jessica Young, Tracey Young, and Kasee Metcalf for always being there for me. But above all I am most grateful for the constant loving support of my family who have always encouraged me to do my best. iii TABLE OF CONTENTS ACKNOWLEDGMENTS .............................................................................................. ii LIST OF TABLES ........................................................................................................vii LIST OF FIGURES ........................................................................................................ ix ABSTRACT ................................................................................................................... xi CHAPTER I Introduction ......................................................................................................... 1 CHAPTER II Carabidae diversity along an altitudinal gradient in a Peruvian cloud forest (Coleoptera) Published in Zookeys (2011) ABSTRACT ...................................................................................................... 18 INTRODUCTION ............................................................................................. 19 METHODS ........................................................................................................ 22 RESULTS .......................................................................................................... 26 DISCUSSION .................................................................................................... 29 LITERATURE CITED ...................................................................................... 34 APPENDIX ....................................................................................................... 51 CHAPTER III Carabid beetle diversity and community composition as related to altitude and seasonality in Andean forests Accepted for publication in Studies on Neotropical Fauna and Environment (December 2013) iv ABSTRACT ...................................................................................................... 53 INTRODUCTION ............................................................................................. 54 MATERIALS AND METHODS........................................................................ 57 RESULTS .......................................................................................................... 59 DISCUSSION .................................................................................................... 61 REFERENCES .................................................................................................. 67 CHAPTER IV Effect of disturbance and inter-annual variation on carabid beetle assemblages in an Andean cloud forest To be submitted to Journal of Tropical Ecology (December 2013) ABSTRACT ...................................................................................................... 81 INTRODUCTION ............................................................................................. 82 METHODS ........................................................................................................ 86 RESULTS .......................................................................................................... 91 DISCUSSION .................................................................................................... 95 LITERATURE CITED .................................................................................... 101 APPENDIX ..................................................................................................... 126 CHAPTER V Patterns of carabid beetle (Coleoptera: Carabidae) morphology along a Neotropical altitudinal gradient in the Peru Submitted to Ecological Entomology (August 2013) ABSTRACT .................................................................................................... 129 INTRODUCTION ........................................................................................... 131 MATERIAL AND METHODS ........................................................................ 135 v RESULTS ........................................................................................................ 140 DISCUSSION .................................................................................................. 144 CONCLUSIONS ............................................................................................. 150 REFERENCES ................................................................................................ 152 CHAPTER VI Carabid beetle assemblages (Coleoptera: Carabidae) on Andean and Appalachian altitudinal gradients To be submitted to Environmental Entomology (December 2013) ABSTRACT .................................................................................................... 169 INTRODUCTION ........................................................................................... 170 MATERIAL AND METHODS ........................................................................ 173 RESULTS ........................................................................................................ 177 DISCUSSION .................................................................................................. 181 REFERENCES CITED .................................................................................... 187 CHAPTER VII CONCLUSIONS AND FUTURE DIRECTIONS ............................................ 204 APPENDIX ................................................................................................................. 212 CURRICULUM VITAE .............................................................................................. 261 vi LIST OF TABLES Table II-1. Locality data for the five altitudinal sites ...................................................... 41 Table II-2. List of all tribes, genera and morphospecies ................................................. 42 Table II-3. Site specificity by taxa ................................................................................. 43 Table III - 1. Comparison of total number of carabid beetle morphospecies between seasons and altitudinal zones .............................................................................. 72 Table III - 2. Carabid beetle diversity and dominance indices by season and altitude ..... 75 Table IV - 1. Summary of individuals and species collected by altitudinal zone ........... 112 Table IV - 2. Description of altitudinal zones for the disturbed gradient ....................... 113 Table IV