© 2004 by Steven James Presley
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© 2004 by Steven James Presley ACKNOWLEDGMENTS Foremost, I thank my major professor, Dr. Michael Willig, for his continual support, encouragement, criticism, and enthusiasm. Mike provided many and varied opportunities for me to grow as a researcher, thinker, educator, and person; hopefully those opportunities were not wasted. Under his guidance I have become a well-rounded scientist, critical thinker, proficient writer, and capable statistician. I am indebted to my committee, Drs. Don Gettinger, Mark McGinley, Daryl Moorhead, Robert Owen, and Richard Strauss. Each has contributed significantly to my growth as a scientist and this dissertation would be lacking if not for their collective guidance. I also thank many faculty members of Texas Tech University, who have provided guidance and enriched my doctoral experience, including Drs. Ray Jackson, Kent Rylander, Michael San Francisco, Charlie Werth, Gene Wilde, and John Zak. I thank Dr. Michael Dini for helping to develop my skills as an instructor. Many fellow graduate students made my time at Texas Tech enjoyable and productive. Stephen Cox was influential in my early development as a doctoral student; we had many discussions over a well-packed bowl that broadened my outlook of the world and biology. Christopher Bloch has been an invaluable office mate during the course of the analysis and writing of my dissertation, being a patient listener and sounding board for my ideas. In addition, I am indebted to Richard Stevens, Celia López-González, Carl Dick, Joel Brant, Chris Higgins, P. Marcos Gorreson, Ed Sobek, Michael Cramer, Kate Lyons, Michelle Secrest, Diane Hall, Brian Croyle, Javier Alvarez, Jeff Roberts, Don Yee, Carla Guthrie, and Kelly Johnson for their friendship, guidance, and support during various epochs of my doctoral studies. I am indebted to Jacy Lewis for her love, support, and encouragement during a critical time when I had abandoned my studies and may not have returned if not for her insistence; thank you. This research generously was supported by grants from the National Science Foundation (NSF) to R. D. Owen and M. R. Willig entitled “Paraguayan Mammals and Their Ectoparasites: an Intensive Survey in a Temperate-Subtropical Interface” (DEB- ii 9400926, DEB-9741543, DEB-9741134). The Office of the Vice-President for Research and Graduate Studies and Office of Research Services at Texas Tech University (D. Schmidly and R. Sweazy) provided substantial financial support during the course of the project. Additional support was provided by the Texas Tech University Graduate School (Summer Research Awards), Department of Biological Sciences (Summer Research Awards and teaching assistantships), and the American Association of Mammalogists (Grant-in-Aid of Research Awards). The Ministério de Agricultura y Ganadería provided significant financial and logistic support. Personnel of the Museo National de Historia Natural del Paraguay and the Convention for International Trade of Endangered Species of Paraguay provided continual encouragement, support, and cooperation, especially noteworthy was the diversity of support that was graciously provided by A. L. Aquino. The Dirección de Parques Nacionales y Vida Silvestre also contributed significantly to the success of the project. The Department of Biological Sciences (J. Burns, M. San Francisco) and the Museum (R. Baker, R. Monk) of Texas Tech University, the Fundación Moises Bertoni (R. Gauto, A. Van Humbeeck), and the Universidad Católica Nuestra Señora de la Asunción (J. Usher) also provided assistance. For access to their land and provision of accommodations while in the field, I especially thank the many owners and administrators of estancias that we visited, personnel of Yaguareté Forests (P. Mueller) and Refugio Faunístico Atingy (M. Cubas), and especially the Fuerzas Armadas de la Republica del Paraguay. For assistance in the field or with logistic arrangements, I thank Heidi Amarilla, Sylvia Frutos, Steven Mezik, Margarita Mieres, and Guillermo Terol. For their dedication to the project and conscientious efforts in the field, I am extremely grateful to Ismael “Pajarito” Mora and Francisco “Mbaracayá” Pintos. I also owe a very large debt of gratitude to Lucy Aquino and her family for making my time in Paraguay as enriching and rewarding personally as it was professionally. Lastly, I need to thank my family for their love, patience and understanding during the seemingly interminable time required to finish “my paper.” In particular, I thank my mother and father for their sacrifice and insistence that I go to college; hopefully, they feel their sacrifice was rewarded sufficiently. iii TABLE OF CONTENTS ACKNOWLEDGMENTS. ii LIST OF TABLES. x LIST OF FIGURES. .xvi CHAPTER I. INTRODUCTION. 1 Literature Cited. .4 II. NATURAL HISTORY OF ARTHROPODS ECTOPARASITIC ON THE BATS OF PARAGUAY. 5 Introduction. 5 Bats and Their Ectoparasites as Model Systems. 5 Study Area and Bat Fauna . .7 Vegetative Setting. 7 Host Assemblage. 8 Ectoparasite Assemblage. 9 Ectoparasite Natural History-Insecta. 9 Streblidae and Nycteribiidae. 10 Polyctenidae. 11 Ischnopsyllidae. .11 Ectoparasite Natural History-Acarina. 11 Spinturnicidae. 12 iv Macronyssidae. 12 Argasidae. 13 Trombiculidae. 14 Chirodiscidae. 15 Myobiidae. 15 Materials and Methods. 16 Field Methods. .16 Laboratory Work. 17 Results. 23 Ectoparasite Assemblages on the Noctilionidae. 24 Ectoparasite Assemblages on the Phyllostomidae. 24 Ectoparasite Assemblages on the Natalidae. .26 Ectoparasite Assemblages on the Vespertilionidae . .26 Ectoparasite Assemblages on the Molossidae. 27 Host Specificity. 29 Ectoparasite Species Abundance Distributions . 30 Discussion. .30 Host-Parasite Associations and Infestation Levels. 30 Host Specificity. 37 v Ectoparasites and Safe Spaces. 39 Streblids and Safe Spaces on Noctilio. .42 Resource Partitioning on Molossids. .44 Implications of Host Phylogeny Based on Ectoparasites. .44 Subfamilial Status of Sturnira. 45 Relationships Among the Vampires, Glossophaga and Carollia. 46 Species Abundance Distributions and Resource Partitioning. 46 Of Birds and Bats. 49 Conclusions. 52 Literature Cited. 54 III. FLYING ISLANDS I: THE EFFECT OF HOST BODY SIZE ON ECTOPARASITE ASSEMBLAGE BIODIVERSITY. 140 Introduction. 140 Bats and Their Ectoparasites as Model Systems. 142 A Host Is an Island, Entire of Itself. 145 Limiting Factors. 145 Materials and Methods. 146 Field Methods. .146 vi Host and Parasite Systematics. 147 Statistical Methods. 148 Analysis of Ectoparasite Abundance. 148 Analysis of Ectoparasite Biodiversity. 149 Results. 152 Total Ectoparasite Abundance. 152 Familial Ectoparasite Abundance. .152 ANCOVA Results for FEA. 154 Specific Ectoparasite Abundance. .154 ANCOVA Results for SEA. 155 Ectoparasite Biodiversity. 155 Discussion. 156 Methodological Considerations. 156 Effects of Host Size on Ectoparasite Biodiversity. 157 Effects of Host Ecology on Ectoparasite Biodiversity. 158 Species Rich, Species Poor. 163 Conclusions. 165 Literature Cited. 167 IV. FLYING ISLANDS II: THE EFFECT OF HOST ABUNDANCE ON ECTOPARASITE ASSEMBLAGE BIODIVERSITY. 199 vii Introduction. 199 Bats and Their Ectoparasites as Model Systems. 201 A Host Is an Island, Entire of Itself. 204 Limiting Factors. 204 Materials and Methods. 207 Field Methods. .207 Host and Parasite Systematics. 207 Statistical Methods. 208 Results. 212 Regional Host Abundance. 212.