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Note to Users NOTE TO USERS This reproduction is the best copy available. UMI LIFE HISTORYAND SEXUALSELECTION Patrick D. Lorch University of Toronto at Mississauga A thesis submittecl in conformity with the recpirements for the degree of Doctor of Philosophy Graduate Department of Zoology University of Toronto Copyright @ 3000 by Patrick D. Lorch University of Toronto at Mississauga l 'except Chapter 5. copyrighted by Society lor Sysdemotic Biology. Reproduced with permission (see p. 123). National Library Bibîioth$uo nationab du Cana a Acquisitions and Acquisitions et Bibliographk Sewices services bibliogrâphiques 395 WdiingtOri SM 385, ni4 Wellington OItawaON K1AONS ûHawaON K1AW canada canada The author has granted a non- L'auteur a accordé une licence non exclusive Licence allowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or seiî reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfichelfilm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substaatial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Abstract Life History and Sexual Selection Patrick D. Lorch University of Toronto at Mississauga Doctor of P hilosophy Graduate Department of Zoology University of Toronto 2000 Three of my thesis chapters use the fecundity by numbers of mates regression slope (Bate- man slopes) to understand how life history allocation patterns can influence the strength of sexual select ion. Males may typically experience st ronger sexual selection because the Bateman slope is Iarger for males than for females. In fact, the ratio of male over female slopes is a way of yuantifying sexual conflict over mating frequency. This conflict may explain the typically higher male motivation to remate relative to females. as well as most other sexual differences (e.g.. shoiviness of peacocks). Animals can allocate energy to different aspects of their life history. and how this allocation affects the strmgth of sexual selection is the central question of most of my thesis. Chapter 2 develops a mode1 for hou sexual selection can be affected by particular life history allocation patterns. 1 focus on reversais in the strength of sexual selection (e.g. females compete for mates), using Bateman slopes for translating changes in allocation to mating effort into changes in the strength of sexuai select ion. My third and fourth empirical chapters test predictions (from Chapter 2) about how the upper limit on sexud selection differs for males and females. Chapter 3 quantifies the effects of a life history t rade-off (between having functional wings and fecundity) on sexual differences in these upper limits. Chapter 4 tests predictions about the relation- ship between the male and female upper limits and the potential for sexual conflict over mating frequency. This chapter also examines why the male upper limit is not greater than the female upper limit as expected. The last two chapters of my thesis are theoretical and do not involve Bateman slopes. Chapter 5 estimates the statistical power of a test designed to ask whether two traits are evolving in a correlated way across a phylogenetic tree. Chapter 6 contains an analytical mode1 of the forces that can cause sexual selection on females to increase. The mode1 examines how the potential sterility of male mates. or less costly risks like mates with ser-ratio distorters. can cause sexual selection for multiple mating in females. Dedicat ion 1 dedicate this thesis to our first child, coming soon, as if to give me something to do when the thesis is done. Acknowledgement s Rather than repeat acknowledgements found in each of the chapters, I will use this space to thank my parents and my partner, and to acknowledge the role of my CO-authorsand rny supervisor. Without the tremendous faith, encouragement and support of my parents I would not have been able to make it this far in what must seem to them an obscure and arcane profession. My wife. Andrea Case. has providd me wit h inspiration. corn fort and all kinds of support. She is the light of my life, the rose of my hcart . While working as a summer lab assistant. Luc Bussière conducted the matings in- volved in Chapter 3 along with entering most of the data. He went on to write the methods of that chapter and to critique drafts of it. Darryl Gwynne suggested the ex- perimeots that led to Chapter 3 and played a major role in designing them. He also guided rny thinking and writing for this and al1 the other chapters of this thesis. 1 am indebted to bot h Luc and Darryl for their tirne and enthusiasm. In the two remaining co-authored chapters. I did al1 the simulations and al1 the initial writing. John Eadie (in Chapter 5. which appeared in a recent issue of Systematic Biology) kept me from stepping too far out on a limb by applying his experience in using comparative phylogenetic tests to do behavioral ecology. He also devised the way we simulated perfect co-evolution and reigned in my unwieldy prose. Lin Chao (in Chapter 6. submitted to .-Lmerican .Vatu- ralist) realized that multiple rnating in fernales would only evolve under a limited set of conditions and suggested 1 work out exactly what the conditions were. He then came up wit h t hr Taylor expansion as an improvernent on my numerical solution to the problem (though taking the partial derivative of this approximation was my idea). 1 would also like to thank both Lin and Xick Collins for encouraging me to build models. Finally. 1 would especially like to thank Darryl. everyone else at Erindale and Locke Rowe for providing such a wonderful atmosphere in which to work. Chapter 2 has been subrnitted to Arnerican .Vaturalist. Contents 1 General introduction 2 Role reversal and Bateman's principle . 5 2.L hbstract .................................... 6 - 2 [ntroduction .................................. 1 2.3 Themodel ................................... 15 2.3.1 SIating ................................. I5 2.3.2 Reproductive energy and fecundity ................. 17 2.3.3 Gift values and sperm allocation ................... 19 2.3.4 Uating distributions ......................... 21 2.3. Sexual select ion gradients ...................... *Pl-- 2.3.6 Error in previous analyt ical results ................. 25 2.4 Results ..................................... '16 2.4.1 Upper limit on sexual select ion ................... 26 2.4.2 Mode1 resuits ............................. 30 5 Discussion ................................... :38 1 Simmons and Parker ......................... 4% 2-52 Arnold and Duval1 .......................... 45 2.5.3 Futurework .............................. 50 4 Summary ............................... 51 2.6 Acknowledgments ............................... 5% 2.7 Appendix ................................... 53 3 Wing-dimorphism and the upper limit of sexual selection. 54 3.1 Abstract .................................... 54 3.2 Introduction .................................. 57 3.2.1 Backqoiind .............................. fi2 2.2 Predict ions .............................. 65 3.3 Materiais and Methods ............................ 66 3.3.1 Collection and rearing ........................ 66 3.3.2 Matings and courtship observations ................. 68 3.3.3 Egg counting. egg laying rates and hatching siiccess ........ 70 3.3.4 Measuring the upper limit on sexual selection ........... 71 3.3.5 Sperm transfer during copulation .................. 72 3.3.6 Statistics ................................ 73 3 .4 Results ..................................... 74 3.4.1 Upper iimits on sexual selection for females and males ...... 74 - c. 3.42 Hatching success ........................... 4, 3.1.3 Courtshipeffects ........................... 79 3.5 Discussion ................................... SO 3.5.1 Upper lirnits on sexual selection for females and males ...... 80 3.5.2 Courtship effects ........................... 83 3.6 Conclusions .................................. 86 3.7 Acknowledgements .............................. $7 4 Potential for sexual conflict over mating frequency. 89 1.1 Abstract .................................... 89 4.2 Introduction .................................. 91 vii 4.3 Methods .................................... 98 1.3.1 Rearing ................................ 98 4.3.2 Datacollection ............................ 100 1.3.3 Analysis ................................ 102 3.4 Results ..................................... 101 4.1.1 Fernale body size ........................... 107 4.4.2 Upper limits on sexual selection ................... 108 4.4.3 Spermatophore weight ........................ 109 1.4.1 Failure of sperm transfer ....................... II2 4.5 Discussion ................................... 11-4 4.6 hcknowledgements .............................. 1'11 5 Power of the concentrated-changes test . 122 5.1 Permissions .................................. 123 5.2 Abstract .................................... 124 5.3 introduction .................................. 125 5.4 Methods .................................... 132 5.5 Results ..................................... 143 5.5. 1 The effect
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