Postcopulatory Sexual Selection in the Soldier Fly Merosargus Cingulatus
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POSTCOPULATORY SEXUAL SELECTION IN THE SOLDIER FLY MEROSARGUS CINGULATUS A dissertation presented to The Faculty of the Graduate School University of Missouri In Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY by FLAVIA BARBOSA Dr. Reginald Cocroft and Dr. H. Carl Gerhardt, Dissertation Supervisors DECEMBER 2011 The undersigned, appointed by the dean of the Graduate School, have examined the dissertation entitled POSTCOPULATORY SEXUAL SELECTION IN THE SOLDIER FLY MEROSARGUS CINGULATUS Presented by Flavia Araújo Barbosa A candidate for the degree of Doctor of Philosophy And hereby certify that, in their opinion, it is worthy of acceptance. Reginald Cocroft, Ph.D. H. Carl Gerhardt, Ph.D. Lori Eggert, , Ph.D. Robert Sites, Ph.D. William Eberhard, Ph.D. ACKNOWLEDGEMENTS This work could not have been accomplished without the support of many, many people. I am thankful for all the advice and support of my supervisors, Rex Cocroft and Carl Gerhardt. They did an amazing job fostering my development as an independent researcher. Not only have they taught me so many necessary skills for being a successful scientist, but they also set an inspiring example. I am also thankful to my dissertation committee, Lori Eggert, Bob Sites, and especially Bill Eberhard. From the beginning of my graduate studies, Bill played a major role in my career, and I owe my success to his advice and encouragement. The days I spent conducting experiments on his compost pile at his backyard in Gamboa are one of my fondest memories of graduate school. I have loved science from an early age, and was lucky to have had wonderful mentors who guided me towards this career. I am grateful to Airton Carrião, for teaching me how to be a scientist. Élida Rabelo, my first research mentor, was extremely influential in my life. I will be forever grateful for all the opportunities and guidance she gave me, as well as for her friendship. I thank Andrea Aspbury, Ray Engeszer and Mike Ryan for their encouragement and help. I am also deeply thankful for Júlio Fontenelle, my undergraduate thesis advisor. Júlio introduced me to my study system, the soldier flies, but he did so much more than that. He taught me invaluable skills and instilled me with a passion for animal behavior. I will be forever grateful to him. ii I started my graduate studies at the University of Texas, and I am grateful to Mike Ryan for being my mentor. I was also lucky to have the opportunity to work alongside a number of talented graduate students, undergraduate researchers and postdocs. I am thankful to all the members of the Ryan lab at the University of Texas, the Gerhardt lab and the Cocroft lab, as well as several fellow graduate students who offered me their support and friendship. Karin Akre, Natalia Biani, Carlos Guarnizo, Katy Klymus, Ashley Groves, Peter Marting, Jeremy Gibson and many, many others, thank you for all the help and all the fun moments. I am especially thankful to Abbie Green, Ximena Bernal and Jen Hamel. The Division of Biological Sciences has so many people who also contributed to my degree. I thank all the wonderful staff for all their help and support, but especially Shannon Dennis, Nila Emerich, Alan Marshall, Josh Hartley and Tyeece Little. I am also grateful to Richard Daniel for the opportunity to teach Bio 1500 for several semesters and for all the fun at the Herpetology field trips. I thank the staff at the Smithsonian Tropical Research Institute, the Organization for Tropical Studies, and Guido Berguido. The governments of Panama and Costa Rica provided research and export permits for this work, and a number of organizations provided financial support. This work could not have been made without the love and support of my family and friends. Para minha mãe, obrigada por uma vida inteira de amor incondicional. Por toda a minha vida, você fez o possível e o impossível para me ajudar a conquistar meus sonhos, e eu não estaria aqui se não fosse por você. Eu também sou gratas às minhas iii irmãs maravilhosas. Joyce, obrigada por uma vida inteira de amizade, e pelas conversas quase diárias durante todo o meu doutorado. Janaina e Vitória, obrigada por fazer minha vida mais feliz. I am thankful for all my friends, especially Rafaela, Leticia and Aninha. I am also immensely grateful for my American family. They have generously welcomed me into their family, and thanks to them, I feel like this country is also my home. Thank you Mary Lou, Walt, Rebecca, Elizabeth, Grandma Gerstle, Grandma and Grandpa Reichert. Finally and most importantly, I am fortunate that my graduate career led me to finding the person with whom I share my life. To Michael, there are no words to thank you for being part of everything I do. Thank you for the love, the encouragement, the constant companionship. Thank you for revising countless versions of the same manuscript and for all the statistical consulting. You had an immense influence in this dissertation in so many ways. Thank you for being my partner in life. iv TABLE OF CONTENTS Acknowledgements ..............................................................................................................ii List of figures ..................................................................................................................... vii List of tables ..................................................................................................................... viii Abstract ............................................................................................................................... ix CHAPTERS 1. Introduction ..…..…….………………………………………………….……………………………………..01 Sperm competition……………………………………………………………………………………………02 Cryptic female choice……………………………………………………………………………………….05 Cryptic male choice………………………………………………………………………………………….07 Study system…………………………………………………………………………………….………………08 Figures………………………………………………………………………………………………………………11 References……………………………………………………………………………………………….……….13 2. Cryptic female choice by female control of oviposition timing in a soldier fly .... 21 Introduction…………………………………………………………………………………………..…………22 Methods………………………………………………………………………………………………………..…24 Results…………………………………………………………………………………………………………..…29 Discussion……………………………………………………………….…………………………………….…30 Tables………………………………………………………………………………..………………………….…34 Figures…………………………………………………………………………………………………………..…36 References………………………………..…………………………………………………………………..…37 v 3. Copulation duration in the soldier fly: the roles of cryptic male choice and sperm competition risk ………………………………….………………………………………………….……….……40 Introduction………………………………………………………………………………………..……………41 Methods…………………………………………………………………………………………..………………45 Results……………………………………………………………………………………………..………………47 Discussion……………………………………………………………….……………………….………………48 Figures………………………………………………………………………………………….….………………52 References………………………………..……………………………………………………..………………56 4. Males responding to sperm competition cues have higher fertilization success in a soldier fly …………………….……………………………………………………………………….……….….…61 Introduction………………………………………………………………………………………..……………62 Methods……………………………………………………………………………………………..……………65 Results…………………………………………………………………………………………………………..…69 Discussion……………………………………………………………….…………………………………….…71 Tables………………………………………………………………………………..………………………….…75 Figures…………………………………………………………………………………………………………..…77 References………………………………..…………………………………………………………………..…80 5. Discussion ................................................................................................................ 85 Summary of results……………………………………………………………………………………..……85 Future directions …………………………………………………………………………………………..…87 References………………………………..………………………………………………………………………90 Vita .................................................................................................................................... 93 vi LIST OF FIGURES CHAPTER 1 Figure 1…………………………………………………………………………………………………………….11 Figure 2…………………………………………………………………………………………………………….12 CHAPTER 2 Figure 1…………………………………………………………………………………………………………….36 CHAPTER 3 Figure 1…………………………………………………………………………………………………………….52 Figure 2……………………………………………………………………………………………..……………..53 Figure 3…………………………………………………………………………………..………………………..54 Figure 4…………………………………………………………………………………………………………….55 CHAPTER 4 Figure 1…………………………………………………………………………………………………………….77 Figure 2…………………………………………………………………………………………………………….78 Figure 3……………………………………………………………………………………..……………………..79 vii LIST OF TABLES CHAPTER 2 Table 1…………….……………………………………………………………………………………………..34 CHAPTER 4 Table 1…………….……………………………………………………………………………………………….75 Table 2…………….……………………………………………………………………………………………….76 viii ABSTRACT My dissertation research focuses on the largely understudied field of postcopulatory (or “cryptic”) mate choice. For the past few decades, it has become increasingly clear that both males and females can perform mate choice during and after copulation, but the implications of this for sexual selection and evolution have been little explored. As part of my dissertation research, I have developed a novel model system for the study of postcopulatory mate choice, the soldier fly Merosargus cingulatus (Diptera: Stratiomyidae). The soldier flies have proven to be an excellent study organism, since they are extremely tractable for field experiments, which I conducted at the Smithsonian Tropical Research Institute field station in Gamboa, Panama, and at the Organization for Tropical Studies field station in La Selva, Costa Rica. The vast majority of studies of postcopulatory