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Information to Users INFORMATION TO USERS This manuscript has been reproduced trom 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 others may be trom any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. ln the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize matarials (a.g., maps, drawings, charts) are reproduœd by sedioning the original, beginning at the upper left-hand corner and continuing tram left to right in equal sedians with small averlaps. 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Bell &Howell Information and Leaming 300 North Zeeb Raad, Ann Arbor, MI 48106-1346 USA 800-521-0600 tvfiCROSATELLITE DNA ANALYSIS OF THE MATING SYSTEM DURING THE FIRST BREEDING PERIOD Of THE FEMALE SNOW CRAB CH/ONOECETES OP/LlO (BRACHYURA, MAJIDAE) Nicola Urbani Department ofAnimal Science Macdonald Campus ofMcGill University Montreal, Canada June 1998 A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfilment ofthe requirements ofthe degree of Doctor ofPhilosophy () Nicola Urbani 1998 National Library Bibliothèque nationale 1+1 of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. rue Wellington Ottawa ON Kl A ON4 Ottawa ON Kl A ON4 canada Canada Your fi,. Votr. reffirencs Our file Notr. r8fsrMCS The author has granted a non­ L'auteur a accordé une licence non exclusive licence allowing the exclusive pennettant 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 fonnats. la fonne de microfiche/film, de reproduction sur papier ou sur format 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 from 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 penmSSlon. autorisation. 0-612-44613-1 Canad~ Suggested short title: Microsatellite DNA Analysis ofthe Snow Crab Mating System. ii ABSTRACT In order to study sperm competition and mating dynamics in the snow crab Chionoecetes opi/io, a genomic library was established with the goal of identifying highly polymorphic microsatellite markers. Six pairs of DNA primers were designed to amplify markers Cop3-4, Cop4-I, CopS, Cop 10, Cop24-3 and Cop 111 by the polymerase chain reaction (peR). AlI markers produced patterns as expected from single loci inherited in a mendelian fashion, except for CapS which revealed a multi-Iocus banding pattern. The cross-amplification ofthe six loci in seven additional crabs species revealed DNA polymorphisms at one or more loci for each species. Markers Cop3-4 and Cop24-3 were used to detennine patemity oflarvae ofprimiparous females bath from the wild and from multiple mating experiments under laboratory settings. The two markers were also used to genotype the contents offemale spennathecae in arder ta detennine the number ofmale genotyes present. Spermathecal contents ofwild-caught females were eut into severa! cross-sections and each section genotyped individually. Histological analysis of spennathecae was canied out to complement genetic data in arder ta elucidate patterns ofsperm competition. Single patemity was observed for the progeny of a1l females. The analysis of laboratory females showed that spenn displacement was the mecbanism by which single patemity was obtained by the last males to mate. The analysis ofwild females revealed that their spermathecae contained on average the spenn of at least 3.7 males. Larvae appeared to he sired by males whose genotypes were found in the spennathecal cross-sections toward the blind-end ofthe spennathecae. This suggested that they were the tirst males to mate with females they guarded until oviposition, and females remated with other males thereafter. Also, a comprehensive account ofthe mating dynamics was carried out in a wild ili population of the Northwest Gulf of Saint Lawrence (Eastern Canada) and demonstrated the existence of intricate male mating hierarchies and assortative pairing processes based on the size and quality ofmates, in a context ofsorne times intense male sexual competition. For the fIlSt time in brachyurans data suggested that female handicaps and secondary sexual characters also influence pair formation. In arder to complement the data on the mating behavior, microsatellite analysis at loci Cop3-4 and Cop24-3 was canied out in five mating pairs involving males grasping primiparous females with their egg clutches. Genetic analysis showed that females had mated on average with 3.4 males, but none ofthe graspers had fathered their mates' eggs. It appeared that these males had smaller shells or shells in poorer conditions than dominant males mating with recentIy molted pubescent females. iv RÉsUMÉ Dans le but d'étudier les stratégies de reproduction et les modalités de competition de spenne chez le crabe des neiges Chionoecetes opi/io, une librairie d'ADN génomique a été élaborée afm d'identifier des marqueurs hypervariables d'ADN microsatellite. Six paires d'amorces ont été construites afin d'amplifier les marqueurs Cop3-4, Cop4-I, Cop5, Cop 10, Cop24-3 et CoplI! par la réaction de polymérisation en chaîne (PCR). Alors que cinq marqueurs révelaient des loci individuels avec des allèles transmis de façon mendel1ienne, le marqueur Cop5 a révelé des patrons multi-Iocus. La présence des six loci a été évaluée chez sept autres espèces de crabe et des polymorphismes ont été détectés à un ou plusieurs loci pour chacune des espèces. Les marqueurs Cop3-4 et Cop24-3 ont ensuite été utilisés afm de déterminer la paternité des larves provenant de femelles primipares sauvages et de femelles primipares utilisées dans des expériences d'accouplements en laboratoire. Les deux marqueurs ont aussi été utilisés pour analyser le contenu des spennathèques de femelles afm de détenniner le nombre de génotypes mâles qu'elles contenaient. Dans le cas des femelles sauvages, le contenu de chaque spermathèque a été coupé en plusieurs sections et le génotype de chacune d'elles a été détenniné individuellement. De plus, une analyse histologique a été effectuée dans le but de compléter les données génétiques afin d'élucider les modalités de competition de sPenne. L'analyse génétique a démontré qu'il Yavait paternité unique pour la progéniture de toutes les femelles. L'analyse des femelles de laboratoire a démontré que le mécanisme de competition de sperme responsable poW' la paternité unique était le déplacement d'éjaculats précedents par celui du dernier mâle accouplé. L'analyse de femelles sauvages a démontré que leur spermathèques contenaient en moyenne le sperme de 3.1 mâles. v Cependan4 les larves de femelles sauvages avaient été fertilisées par les mâles dont le génotype apparaissait dans les sections des spennathèques les plus reculées. Ceci suggère qu'ils étaient les premiers a s'accoupler avec des femelles qu'ils auraient défendues jusqu'à oviposition et que les femelles se seraient réaccouplees par la suite. De plus, les dynamiques d'accouplemet chez une population du nord-ouest du Golfe du Saint-Laurent dans l'est du Canada ont été analysées. Les résultats ont démontré l'existance d'une hiérachie d'accouplement chez les mâles, et de processus de formation de couples qui sembleraient se baser sur la taille et la qualité des partenaires dans un contexte de compétition entre mâles parfois intense. L'analyse a également montré pour la premiere fois au sein des Brachyura que des caractères sexuels secondaires chez les femelles peuvent aussi influencer la formation de couples. Afin de complémenter les données sur les dynamiques d'accouplemen4 une analyse génétique à partir des loci Cop3-4 et Cop24..3 a été éffectuée chez cinq couples impliquant des mâles saisissant des femelles primipares avec leur portées d'oeufs. L'analyse a démontré que les femelles s'étaient accouplées en moyenne avec 3.4 mâles et que les mâles saisissant les femelles n'étaient pas les pères de leurs oeufs. Les cinq mâles analysés étaient plus Petit ou dans des conditions moins bonnes que les mâles dominants s'accouplant avec des femelles pubères ayant reçemment mué. vi Questa tesi e dedicata a papofiglio Uberto Urbani. vü ACKNOWLEDGMENTS There are many people l wish to thank for their help in completing this work. First, l would like to sincerely thank Dr. Urs KuhnIein, my supervisor, for his guidance and patience throughout my studies. l am grateful for the freedom he gave me and the Many opportunities ta go out on croises and ta attend scientific meetings. Many thanks to Ors. Bernard Sainte-Marie, Jean-Marie Sévigny, Samuel Aggrey and David Zadwomy for their guidance and numerous discussions. [ wish to thank Ors. Bernard Sainte-Marie and Jean-Marie Sévigny as weil as Éric Parent for their friendship and hospitality during my stays at the Maurice Lamontagne Instltute in Mont-Joli. Many thanks to the crew ofthe CaJanusII for being so welcoming and so nice when l was sea...sick.
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