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INFORMATION TO USERS This manuscript bas been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, sorne thesis and dissertation copies are in typewriter face, while athers may be trom any type ofcomputer printer. The quality ofthis reproduction is dependent upon the quality ofthe copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper aIignment cao adversely affect reproduction. In the unlikely event that the author did not send UMl a complete manuscript and there are missing pages, these will be noted. AIso, if unauthorized copyright materia! had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are repradueed by sectioning the origin~ beginning at the upper left-hand corner and continuing trom left ta right in equai sections with small overlaps. Each original is a1so photographed in one exp0 sure and is included in reduced form at the back ofthe book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographie prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to arder. UMI A Bell et Howell Information Company 300 North Zeeb Roa~ Ann Arbor MI 48106-1346 USA 313n61-4700 800/521-0600 SEXUAL SIZE DIMORPHISM AND SELECTION IN THE WATERSTRJDER AQUARJUS REltflGIS Richard F. Preziosi Department of Biology, MeGill University, Montreal A thesis submitted to the Faeulty of Graduate Studies and Research in partial fulfillment of the requirements for the degree Doelor of Philosophy January 1997 © Richard F. Preziosi r... National Ubrary Bibliothèque nationale 1+1 of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. rue Wellington Ottawa ON K1 A ON4 Ottawa ON K1 A QN4 Canada canada The author bas granted a non­ L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Library ofCanada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies ofthis thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/film, de reproduction sur papier ou sur fonnat électronique. The author retains ownership ofthe L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts frOID it Ni la thèse ni des extraits substantiels may be prînted or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son pemusslon. autorisation. 0-612-30363-2 Canadrl Abstract Sexual size dimorphism (S5D), or the difference in body size between males and females, is common in almost aH taxa of animaIs and is usually assumed to be adaptive. Darwin's two main mechanisms for the evolution of SSO. sexual selection for larger males and fecundity selection for larger females. have often been demonstrated. However, males and females frequently share both genes and environment and more recent papers have noted that males and females must experience differences in lifetime selection on body size for SSD to be maintained. Over two generations l examined lifetime selection acting on adult body size (totallength) in a common insect where females are larger than males, the waterstrider Aquarius remigis. Both fecundity selection for larger females and sexual selection for larger males are oeeurring in this speeies and both selective forces appear to target specifie components of body size rather than total length~ sexual selection targeting male genitallength and fecundity selection targeting female abdomen length. While body size did .'lot appear to influence adult prereproductive survival. longevity during the reproductive season was negatively related to body size for both sexes. When the opposing selection of reproductive success and reproductive longevity are combined. bath males and females have an intennediate optimum body size. A remarkable result of this stabilizing selection was that the optimum size of males was smaller than that of females. 1also examined the repeatability of reproductive success in both sexes and the trade-off between egg size and egg number. Finally. estimates of the quantitative genetic basis of the traits examined indicate that both male and female body size. and components of body size. are heritable and can respond to the selection detected. Components of body size in A. remigis are variable in bom the degree and direction of sexual dimorphism and the genetic analysis indicates partial isolation of dimorphic and monomorphic traits within each sex. Il { Résumé Le dimorphisme sexuel de la taille (DST). ou la différence entre la taille des mâles et des femelles. est commun dans la majorité des taxons du règne animal. où il est généralement considéré adaptatif. Les deux mécanismes proposés par Darwin pour expliquer l'évolution du DST. soit la sélection sexuelle qui favorise les plus grands mâles et la sélection sur la fécondité qui avantage les grandes femelles. ont souvent été démontrés. Cependant. les mâles et les femelles partagent souvent leurs gènes et environnement et il a été noté récemment qu'ils devraient être exposés à des pressions de sélection divergentes sur la taille pour que le DST se maintienne. J'ai étudié sur deux générations la sélection qui affectait la taille adulte (longueur totale) de l'insecte Aquarius remigis, une espèce commune de patineuse où la femelle est plus grosse que le mâle. Les deux types de sélection proposés par Darwin ont été détectés chez cette espèce et les pressions de sélection semblent agir spécifiquement sur certaines composantes de la taille au lieu d'affecter la longueur totale; la sélection sexuelle agit sur la longueur du génitaIia des mâles et la sélection sur la fécondité affecte la longueur de l'abdomen des femelles. Alors que la taille ne semblait pas influencer la survie des adults préreproductifs. une relation inverse entre la longévité durant la saison de reproduction et la taille a été détectée chez les deux sexes. Les mâles et les femelles démontrent une taille optimale intermédiaire lorsque les pressions de sélection inverses qui agissent durant la saison de reproduction sur le succès reproducteur et la longévité sont combinées. Un résultat remarquable est que la taille optimale prédite par cette sélection stabilisatrice est effectivement plus petite chez les mâles que chez les femelles. J'ai égaiement mesuré la reproductibilité des comportements qui affectent le succès reproducteur chez les deux sexes et le compromis entre la taille des oeufs et la fécondité. Finalenlent, une étude de génétique quantitative suggère que la taille ainsi que les caractères qui contribuent à la taille sont héréditaires chez les deux sexes et donc qu'ils pourraient répondre aux pressions de sélection détectées. Les caractères qui contribuent à la taille chez A. remigis démontrent différents degrés et directions de dimorphisme sexuel, alors que l'étude génétique révèle que les traits monomorphiques et dimorphiques sont partiellement isolés ".r dans chaque sexe. III • Table of Contents Abstract i Résumé ii Table of Contents iii List of Tables v List of Figures vi Preface vii REMARKS ON STYLE AND AUTHORSHIP vii Original Contributions to Knowledge ix Acknowledgements xii General Introduction 1 Chapter 1. Sexual size dimorphism and selection in the wild in the waterstrider Aquarius remigis: Body size. components of body size and male mating success 15 ASSTRACT ............................................................................................................................•.................... 16 [NTRODUCTION 17 METHODS 20 REsULTS 26 DISCUSSION .......................................................................•....................................................................... 29 :\CI<J"IlOWLEDG~fENTS 34 LITERATURE CITED ...........••....•............................................................•...•................................................. 35 Chapter 2: Body size and fecundity in the waterstrider Aquarius remigis: A test of Darwin's fecundity advantage hypothesis , 51 ASSTRACT .,................................•.........................................................•..........................................•......... 52 [r-oTRODUCTION .................•..............................................................................................•.....•................... 53 METHODS 57 REsULTS 60 DISCUSSION .............................•...•........................,...............................•.............................•..•................... 62 AcKNOWLEDGMENTS.•.......................................................................................................•...•......•............ 65 REFERENCES ....................................................................................................•...•.....................................66 Chapter 3: Sexual size dimorphism and selection in the wild in the waterstrider Aquarius remigis: lifetime fecundity selection on female total length and its components 78 ASSTRACT .,........................•......................................................•..............•...............................................
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