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Good Genes Or Sexy Sons? Testing the Benefits of Female GOOD GENES OR SEXY SONS? TESTING THE BENEFITS OF FEMALE MATE CHOICE IN THE PAiNTED TURTLE, CHRYSEMYS PICTA A Thesis Presented to The Faculty of Graduate Studies Of The University of Guelph In partid fulfiiiment of requiremtns for the degree of Master of Science August, 2000 National Library Bibiiithèque nafinale 1*1 .Cam, du Canada Acquisiîions and Acquisitions et Bibliographie Services sefvices bibliographiques 345 Wellinglon Street m. rue welrigton Ottawa ON K1A ON4 CNmwaON KlAM CaMda Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence dowing the exclusive permettant à la National Library of Canada to Bbiiotheque nationale du Canada de reproduce, loan, distniute or seil reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/6llm, 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 substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or othewise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. GOOD GENES OR SEXY SONS? TESTING THE BENEFITS OF FEMALE MATE CHOiCE IN THE PAINTED TURTLE, CHRYSEMYS PICTA Seanna J. McTaggart Advisor: University of Guelph, 2000 Professor R.J. Brooks The "good-genes" and the "sexy-sonJ1hypotheses explain the evolution of Eemale preferences in species where femaies do not receive direct benefits from mating. The good genes hypothesis predicts that femdes prefer certain males because they confer a genetic benefit to her offspring, whereas the sexy son hypothesis predicts no such benefit. This study tested these opposing predictions in the painted turtle (Chrysernys picta). Paternity of twenty clutches from a small, closed popdation was determined using microsatellites. Field observation revealed that female choice was active in this species, and molecular data confirrned that mating was non-random. Ciutches of desthat sired more than one clutch had higher hatching frequency than clutches of males only siring one cIutch, supporting the good-genes hypothesis. Furthmore, desof higher reproductive success had smaller carapaces than desof lower reproductive success. ACKNOWLEDGMENTS The support 1 have received during this project has been vital to me and to the completion of this project, and 1 am gratefui to have the opportunity to mite these pages. To those who read this thesis please keep in mind the iremendous effort, time and patience that has been so generously given by each of the following people. For my introduction into the world of herpetology 1 thank every member of the Brooks' Iab: Sara Ashpole, Shane de SoiIa, Sarah Holt, Kevin Judge, Cam MacDonald, Elaine Matthews, Janet Lochead, Jake Rouse, Kim Smith, and Rob WiIIson. i would especidy like to acknowledge Jon Edmonds and Dave Cunnington for their valuable friendships. For an unforgettable and fruitful field season at the Wildlife Research Station in Algonquin Park 1 thank Mattman, Rob van Vlaenderan, Yemisi Dare, Mandy Karsch, and Sara Wicket, and most importantly the turtles at WoiE Howl Pond. I completed my second year supported primarily by three individuais. Teri Crease, John Colbourne and Cheryl kokopwich al1 provided unlimited time and energy into helping me trouble-shoot my way through technical difficulties. More importantly ail three met my frustrations and fears with inhnite patience and by always voicing their faith in me. During the writing of this thesis, 1 have my parents, Anne and Gault, to thank above everyone else. They generously provided me with a peaceful home to mite in, and supported me mesemediy throughout each part of the thesis. 1also must thank the foIlowing hiends who without their emotional support 1 would probably have left this piece of work unfinished: Ryan Gregory, Jen McCourt, Keith McTaggart and Kara Vlasrnan. To al1 of my cornmittee members 1 am grateful for helpful comrnents and advice. Specifically, to Tom Nuddç 1 am grateful for his positive outlook, clarity of thought and his faith in me. Teri Crease 1 thank for providing me with such an inspirational mentor figure. Finally, 1 thank my advisor Ron Brooks for giving me the space to finci my own way, and for providing inspirationai thoughts that reach far beyond the sape of this project. Acknowledgements Table of Contents List of Tables List of Figures Introduction Sexual selection Turtles as study organisms Study population Methods Sarnple collection Tissue sarnples Morphology FI generation DNA extraction Molecular markers DNA amplification Gel electrophoresis and DNA visuaIization Scoring native gels Genotype assignment Data analysis Results Field observations Data analyses Pa ternal assignment Successfd versus unsuccessful males Sexy son versus good genes Factors explaining hatchling DUD scores Discussion Fernale choice Behavioural evidence Morphological evidence Sexy son versus good genes Hatchling mortality Conclusions Tables Figures Literature cited Appendix 1 Table Page 1 Indirect benefits of females in species with no patemai care: 37 studies examining if female mate choice effects offspring viabiiity . 2 Studies uing moIecular markers to track paternity in turties. 3 Sequences of the three microsatellite primers. 4 Potentiai and realized male reproductive success for males siring study clutches. Figure Page Graphitai representation of the predictions of the competing 41 good-genes and sexy-son hypotheses for the evolution of female mate choice. Ge1 depicting a mother and her five hatchlings. 42 Gel showing fdyA19 at 1- Cp2. 43 Gel showing farnily A19 and some putative fathers at locus 44 CplO. Gel showing farnily A19 with putative fathers at locus Cp3. 45 An example of a gel cantaining desamples. 46 Allele frequency distribution for microsatellite Iocus Cp2, 47 frorn a sample of a population of Chrysemys pictu in Wolf Howl Pond, Algonquin Park, Ontario. Plot of midarapace length of desthat did not sire any of 48 the study clutches (Group 0) and those that did (Croups land 2). Mot of average third ciaw length for males with no study 49 clutches sired (Group O) and rnaies that did sire ail study chtches (Groups 1 and 2). Frequency histogram O£ male reproductive success from a 50 sampk of 20 clutches taken hmWolf Howl Pond, Aigonquin Park, Ontario. Plot of days untii death (Dm)scores for hatchIings from 51 Group 1 and Group 2. INTRODUCIION Semial selection To explain the evolution of conspicuous male traits that appear to hinder rather than enhance survival, Darwin (1874) advanced the theory of sexual selection. He postulated that sexuai selection &ove the evolution of these traits because males that bore them gained an advantage over other males in acquiring mates. This idea has been at the heart of a plethora of theoreticai and ernpirical studies. Despite this intense interest, an understanding of the evolution and maintenance of mating patterns remains elusive. Initial studies of sexual seiection focused on determining if male mating success was non-random, if the pattern of mating was correiated with variation in male ornaments, and if the pattern might be caused by female choice (Jennions and Petrie 1997). The number of studies confirming these two patterns with respect to female preferences is large (reviewed in Johnstone 1995, Andersson 1994; Ryan and Keddy-Hector 1992). The next body of research to emerge explored the possible selection pressures that could maintain fdepreferences (Jennions and Petrie 1997). Preferences are any semry and/or behavioud tendency that influence individuals to mate with certain phenotypes (Kirkpatrick, 1997). Mate choice is defined as the actual pattern of mating that arises, and preferences may or may not strongly influence that pattern. There are two primary explanations for the maintenance of preferences: (1) preferences are maintained by direct selection because they increase female survival (Reynolds and Gross 1992). (2) selection indirectly maintains preferences because mating with preferred males either increases the number and/or viabiiity of her offspring (good-genes) or increases the attractiveness of her sons, which increases their abiiity to obtain mates (sexy-sons) (Fisher 1958). Direct and indirect seletion may act either singly or in concert, and must be examined in the context of the species under study in order to determine what influence they rnay have on female preferences. When females do not gain direct benefits from mate choice, other than sperm to fertilize her eggs, male courtship behaviour rnay be the only means by which fernales have the opportunity to assess potential mates. in the absence of direct selection for female choice, Fisher (1958) suggested two hypotheses that might explain how indirect selection could drive the evolution of fde preferences for male ornarnents. Fust, he suggested that male traits are indicators of high heritable quality, which will translate into offspring fitness. This is comrnonly known as the "good-gene hypothesis". Genetic quality in these cases is defined as differential viability or numbers of offspring among different males (Boake 1986, Qutton-Brock 1992). This hypothesis predicts that females mating with more preferred or attractive males will have greater hatchg frequency, and/or any measurement of offspring viabüity compared with femaies with weaker, or no preference (Figure la) (Fisher 1958). This prediction has been tested and supported by a number of studies (Table 1). The "sexy-soni', or runaway selection hypothesis, predicts that the offspring of femaies who have mated with the preferred males, are no more viable or numerous than offspring of non-discriminating females (Figure Zb) (Fisher 1958). Fernale preferences evolve in the absence of increased offspring viability because femaies with a strong preference simply have sons that inherit the preferred trait, causing them to enjoy a higher than average rnating success. The daughters of these females inherit the preference, rnaintaining it in the population.
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