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The Evolutionary Ecology of Sexual Conflict and Condition-Dependence in an Insect Mating System

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The Evolutionary Ecology of Sexual Conflict and Condition-Dependence in an Insect Mating System The evolutionary ecology of sexual conflict and condition-dependence in an insect mating system by Jennifer Christine Perry A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Ecology and Evolutionary Biology University of Toronto © Copyright by Jennifer Christine Perry 2010 The evolutionary ecology of sexual conflict and condition-dependence in an insect mating system Jennifer Christine Perry Doctor of Philosophy Ecology and Evolutionary Biology University of Toronto 2010 Abstract Sexual conflict and condition-dependent trait expression have emerged as major themes in sexual selection. There is now considerable evidence suggesting that both conflict and condition- dependence can drive the evolution and expression of sexual traits; still, important questions remain concerning the extent to which conflict shapes sexual traits and the role of condition in mediating conflict. Here, I address these two themes in studies of a ladybird mating system. One set of studies investigates the function and economics of potentially antagonistic traits – nuptial gifts and female mating resistance – while another examines condition-dependence in mating resistance and male ejaculate composition. Nuptial gifts are often considered beneficial to females, but recent thinking suggests they may also allow males to manipulate females, raising the possibility of conflict. I demonstrate that male ladybirds benefit from nuptial feeding by their mates through reduced re-mating frequency. Benefits to female reproduction and lifespan, however, are weak or non-existent. These results show that although males gain from transferring gifts that influence female behaviour, females experience neither harm nor benefit. I next tested the hypothesis that nuptial feeding is maintained – despite an absence of benefits – because female foraging is generally elevated after mating. However, although ii females indeed display strongly increased foraging after mating, this response did not increase nuptial feeding. Recent studies suggest that individual condition may affect the economics of mating and extent of conflict. Female ladybirds vigorously resist mating, and I show that (1) resistance is condition- dependent, with low-condition females displaying more resistance, and (2) resistance functions to minimize superfluous matings (sexual conflict), rather than to select among males (indirectly benefiting females). Resistance generates selection favouring large males; thus, this work demonstrates that ecological circumstances, through influencing condition, affect the strength of sexual selection. Finally, male condition may influence investment in ejaculate components, but condition- dependence in ejaculate composition is currently poorly understood. I show that, in agreement with theory, males in poor condition transfer smaller ejaculates that nonetheless contain more sperm, but less seminal fluid. Taken together, this work highlights both the value of economic studies in evaluating sexual conflict, and the significance of condition-dependence for sexual selection. iii Acknowledgements I am grateful to those who ably and cheerfully assisted with the experiments presented in this thesis: Alya Ahsan, Christine Heung, Kevin Sha, Diana Sharpe and Crystal Tse. I gratefully acknowledge experimental animals supplied by Suzanne Lommen and Lee Henry. Perhaps the most enjoyable aspect of my doctoral research has been stimulating conversation with colleagues. For this I thank Aneil Agrawal, Jay Biernaskie, Amber Budden, Sean Clark, Asher Cutter, Elah Feder, Darryl Gwynne, Don Jackson, Shannon McCauley, Michael Majerus, David Punzalan, Helen Rodd, Mariana Wolfner and Minyoung Wyman, and above all my advisor Locke Rowe. The manuscripts were vastly improved by feedback from Jay Biernaskie, Asher Cutter, Darryl Gwynne, Lucia Kwan, Tristan Long, David Punzalan, Maxence Salomon and Jessica Ward. Any errors that remain are mine alone. I acknowledge with gratitude the members of my supervising committee, Asher Cutter and Darryl Gwynne, for their generous attention and useful criticism. I also appreciate the thoughtful feedback of the additional members of my appraisal and defence committees: Helen Rodd, Nina Wedell and Deborah McLennan. I thank my advisor Locke Rowe, who has been more than I could have hoped for in a mentor. Thank you for your commitment to making me a better scientist. I was supported financially by a Canada Graduate Scholarship from the Natural Sciences and Engineering Research Council of Canada (NSERC); the Dr. F.M. Hill Ontario Graduate Scholarship in Science and Technology; an Entomological Society of Canada Postgraduate Award; a W. John D. Eberlie Travel Research Grant from the Toronto Entomological Association; and a Ramsay Wright Scholarship provided by the Department of Ecology and Evolutionary Biology at the University of Toronto. My research was also supported by grants to Locke Rowe from NSERC and the Canada Research Chairs program. I thank these agencies. It is a pleasure to warmly thank to my parents, Brian and Sharon Perry, for their support throughout my postgraduate career and all my life. iv Finally I thank my partner Jay Biernaskie. I promise to listen to you carefully, consider your ideas, and be open to the many things I can learn from you. Statement of contributions Chapters 2, 3, 5 and 6 have been published as co-authored papers. All of these papers were co- authored with my senior advisor, Locke Rowe. In each case, we developed the study’s aims and experimental methods in collaboration; I conducted the experiments, analyzed the data and was the primary author of the manuscripts. Chapter 5 was also co-authored with Diana Sharpe. Diana refined the experimental methods and we conducted the experiments together. v Table of Contents Acknowledgments ............................................................................................................................................. iv List of Tables ..................................................................................................................................................... ix List of Figures ..................................................................................................................................................... x Chapter 1 General introduction ....................................................................................................................... 1 1 General introduction .................................................................................................................................... 1 1.1 Sexual conflict ....................................................................................................................................... 1 1.2 Condition ............................................................................................................................................... 3 1.3 Nuptial gifts and Adalia bipunctata ...................................................................................................... 3 1.4 The evolutionary ecology of sexual conflict and condition-dependence in Adalia bipunctata ................................................................................................................................................. 5 Chapter 2 Ingested spermatophores accelerate reproduction and increase mating resistance, but are not a source of sexual conflict ............................................................................................................. 8 2 Abstract .......................................................................................................................................................... 8 2.1 Introduction .......................................................................................................................................... 9 2.2 Methods ............................................................................................................................................... 12 2.3 Results .................................................................................................................................................. 15 2.4 Discussion............................................................................................................................................ 17 2.5 Appendices .......................................................................................................................................... 22 2.6 Figures .................................................................................................................................................. 24 2.7 Tables ................................................................................................................................................... 27 Chapter 3 Neither mating rate nor spermatophore feeding influences longevity in a ladybird beetle ............................................................................................................................................................. 28 3 Abstract ........................................................................................................................................................ 28 3.1 Introduction ........................................................................................................................................ 28 3.2 Methods ..............................................................................................................................................
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