It has been almost 150 years since Darwin’s “The Origin of Species” was published, yet the mechanisms that lead to the creation of new species are still a topic of fervent research in Evolutionary Biology. In the last several decades, there has been considerable interest in how genetic drift and adaptive processes internal to populations may lead to population divergence. It has been recognized that intersexual co-evolution may drive the rapid evolution of reproductive traits, leading to reproductive isolation between populations in allopatry. To explore the nature of this process, I conducted a series of interpopulation interaction experiments using a set of laboratory-evolved Drosophila melanogaster populations. These “B” (for baseline) populations share a common ancestor and had evolved in identical habitats in allopatry for 650-700 generations prior to my experiments. When different populations were combined in competitive fitness assays, I found that foreign males usually suffer depressed fitness when competing with males who share a common evolutionary history with the focal females. I selected a subset of B populations to investigate potential sources of variation that could explain this result. I examined; 1) characters involved in post-copulatory interactions between mates and 2) traits involved in mating behaviour. I found that sperm precedence patterns varied significantly depending on female population of origin and suggestive evidence that males between populations vary in ability to compete in sperm offence. Males also varied in mating behaviour between populations, however we found no evidence that divergence of these traits was correlated to that of traits involved in post-copulatory interactions, suggesting that understanding how intersexual co-evolution drives population divergence requires considering independent components of the reproductive phenotype and their contributions to overall fitness differences. / Thesis (Master, Biology) -- Queen's University, 2008-08-13 12:29:52.602

Sexual differences are dramatic and widespread across taxa. However, a common genome between males and females should hinder phenotypic divergence. In this thesis I have used experimental, genomic, and theoretical approaches to study processes that can facilitate and maintain differences between males and females. I studied two mechanisms for the evolution of sexual dimorphism - condition-dependence and gene duplication. If sex-specific traits are costly, then individuals should only express such traits when they possess enough resources to do so. I experimentally manipulated adult condition and found that the sex-biased gene expression depends on condition. Second, duplication events can permit different gene copies to adopt sex-specific expression. I showed that half of all duplicate families have paralogs with different sex-biased expression patterns between members. I investigated how current sexual dimorphism may support novel dimorphism. With regards gene duplication, I found that related duplicates did not always have different expression patterns. However, duplicating a pre-existing sex-biased gene effectively increases organismal sexual dimorphism overall. From a theoretical perspective, I investigated how sexually dimorphic recombination rates allow novel sexually antagonistic variation to invade. Male and female recombination rates separately affect invasion probabilities of new alleles. Finally, I examined the assumption that a common genetic architecture impedes the evolution of sexual dimorphism. First, I conducted a literature review to test whether additive genetic variances in shared traits were different between the sexes. There were few significant statistical differences. However, extreme male-biased variances were more common than extreme female-biased variances. Sexual dimorphism is expected to evolve easily in such traits. Second, I compared these results to findings from the multivariate literature. In contrast to single trait studies, almost all multivariate studies of sexual dimorphism have found variance differences, both in magnitude and orientation, between males and females. Overall, this thesis concludes that sexual dimorphism can evolve by processes that generate novel sexual dimorphism or that take advantage of pre-existing dimorphism. Furthermore, a common genome is not necessarily a strong barrier if genetic variances differ between the sexes. It will be an exciting challenge to understand how mutation and selection work together to allow organisms differ in their ability evolve sexual dimorphism.

Variation in condition can result from environmental heterogeneity or genetic variation affecting resource acquisition and processing ability. Although condition should be positively correlated with the fitness of both males and females, the strength of selection on condition may differ between the sexes due to differences in reproductive variance. Sexual selection on male condition has been proposed to reduce mutation load, but only if selection is greater on males than females. To investigate the strength of selection on condition, the quality of the larval environment was used to manipulate the condition of Drosophila melanogaster. Additionally, selection was measured when the availability of key resources for females (live yeast) and males (access to females) were altered. Overall, selection was found to be stronger on males than females. However, selection on males weakened under a female-biased sex ratio, whereas selection on females was not significantly affected by the abundance of live yeast.

Sexual differences are dramatic and widespread across taxa. However, a common genome between males and females should hinder phenotypic divergence. In this thesis I have used experimental, genomic, and theoretical approaches to study processes that can facilitate and maintain differences between males and females. I studied two mechanisms for the evolution of sexual dimorphism - condition-dependence and gene duplication. If sex-specific traits are costly, then individuals should only express such traits when they possess enough resources to do so. I experimentally manipulated adult condition and found that the sex-biased gene expression depends on condition. Second, duplication events can permit different gene copies to adopt sex-specific expression. I showed that half of all duplicate families have paralogs with different sex-biased expression patterns between members. I investigated how current sexual dimorphism may support novel dimorphism. With regards gene duplication, I found that related duplicates did not always have different expression patterns. However, duplicating a pre-existing sex-biased gene effectively increases organismal sexual dimorphism overall. From a theoretical perspective, I investigated how sexually dimorphic recombination rates allow novel sexually antagonistic variation to invade. Male and female recombination rates separately affect invasion probabilities of new alleles. Finally, I examined the assumption that a common genetic architecture impedes the evolution of sexual dimorphism. First, I conducted a literature review to test whether additive genetic variances in shared traits were different between the sexes. There were few significant statistical differences. However, extreme male-biased variances were more common than extreme female-biased variances. Sexual dimorphism is expected to evolve easily in such traits. Second, I compared these results to findings from the multivariate literature. In contrast to single trait studies, almost all multivariate studies of sexual dimorphism have found variance differences, both in magnitude and orientation, between males and females. Overall, this thesis concludes that sexual dimorphism can evolve by processes that generate novel sexual dimorphism or that take advantage of pre-existing dimorphism. Furthermore, a common genome is not necessarily a strong barrier if genetic variances differ between the sexes. It will be an exciting challenge to understand how mutation and selection work together to allow organisms differ in their ability evolve sexual dimorphism.

Variation in condition can result from environmental heterogeneity or genetic variation affecting resource acquisition and processing ability. Although condition should be positively correlated with the fitness of both males and females, the strength of selection on condition may differ between the sexes due to differences in reproductive variance. Sexual selection on male condition has been proposed to reduce mutation load, but only if selection is greater on males than females. To investigate the strength of selection on condition, the quality of the larval environment was used to manipulate the condition of Drosophila melanogaster. Additionally, selection was measured when the availability of key resources for females (live yeast) and males (access to females) were altered. Overall, selection was found to be stronger on males than females. However, selection on males weakened under a female-biased sex ratio, whereas selection on females was not significantly affected by the abundance of live yeast.

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The epaulets of male Red-winged Blackbirds (Agelaius phoeniceus) function in both intersexual and intrasexual contexts, but there is little evidence that they covary with reproductive success in this well-studied species. We used path analysis of male/territory traits, mating success, and reproductive success of unmanipulated males to estimate current directional selection. Territory size had a positive effect on number of within-pair mates. Number of extra-pair mates had a positive effect on number of extra-pair fledglings, and number of within-pair and extra-pair fledglings had positive effects on total number of fledglings. We also reddened epaulets of free-living territorial males to determine whether manipulated and control males differ in territorial behavior, mating success, or reproductive success. Compared with control males, males with reddened epaulets incurred elevated rates of trespassing and territorial challenges, lost their territories more often, and were unable to produce extra-pair offspring. Despite these differences, however, the realized reproductive success of experimental and control males did not differ significantly, perhaps because males with reddened epaulets devoted more time to anti-predator vigilance and were more aggressive toward a simulated predator than were control males. An apparent lack of current sexual selection on epaulet color may be a form of counter-balancing sexual selection in which male aggression against redder epaulets opposes female preference for redder epaulets, or it might be the result of males compensating for lost extra-pair fertilizations by increasing their parental care.

Pendant de nombreuses années, les femelles ont été décrites comme passives dans les interactions mâles-femelles et ont parfois été négligées dans les études sur les comportements reproducteurs et la communication animale. Pourtant, il est de plus en plus évident que les comportements des femelles influencent ceux des mâles et que les mâles peuvent ajuster leurs comportements de parades aux comportements des femelles. L’objectif principal de cette thèse est de mieux comprendre les signaux que les femelles émettent lors d’interactions avec un mâle dans un contexte de reproduction chez le canari domestique. L’ensemble de mes résultats montre tout d’abord que les femelles utilisent des signaux de parades visuels et acoustiques, c’est-à-dire des postures de sollicitation à l’accouplement et des trilles-spécifiques de femelles, comme une invitation à s’accoupler, mais qu’elles peuvent aussi les utiliser pour inciter le mâle à parader et l’aider à échantillonner la qualité de partenaires potentiels. D’autre part, ces deux signaux n’auraient pas la même efficacité selon le contexte d’émission. Ensuite, il semblerait que les signaux véhiculés par la modalité visuelle pourraient jouer un rôle plus important qu’on ne le pensait jusque-là dans les interactions intersexuelles chez cette espèce. Enfin, cette thèse a aussi pu apporter de nouveaux éléments concernant les préférences des femelles pour les chants de mâles et a montré que les méthodes utilisées en laboratoire pour tester les préférences des femelles étaient fiables et congruentes. Cette thèse permet de contribuer aux recherches grandissantes mettant en évidence le rôle important de la femelle dans les interactions intersexuelles. / Much studies on reproduction and animal communication have considered the female as the passive sex; the role of the female during male-female interactions have often been overlooked. However, there is growing evidence that female behaviours can affect those of the males and that males can adjust their courtships to female behaviours. The aim of this thesis is to understand the signals produced by females during interactions with a male in a reproductive context in the domestic canary. Overall, my results show that females not only use their visual and acoustic signals, the copulation solicitation display and the female-specific trills, as an invitation to copulate but also to incite male to sing as an aid to sample potential mates. Then, these two signals could not have the same efficacy in different contexts of transmission. Moreover, the visual components of the communication seem to be more important than previously thought during intersexual interactions in this species. Finally, this thesis provides new elements about the female preferences for male songs and shows that methods used to test female preferences in laboratory are reliable and congruent. This thesis contributes to the growing number of researches showing that females play an active role in intersexual interactions.

Chez les espèces sociales, les individus des deux sexes peuvent interférer avec la sexualité et les alliances sociales des femelles, ce qui peut influencer les stratégies reproductives des femelles. Un regain d'intérêt récent pour l'action de la sélection sexuelle chez les femelles a mis en évidence que la compétition entre femelles pour monopoliser les ressources reproductives, comme les partenaires sexuels ou les soins aux petits, est prévalente. Cependant, jusqu'à présent, la compétition reproductive entre femelles a reçu peu d'attention chez les espèces polygynes. Nous avons donc étudié les déterminants de la compétition reproductive entre femelles dans une société primate polygyne, dans une population naturelle de babouins chacma, en Namibie. Nos résultats montrent que l'agression est plus intense entre les femelles qui sont en synchronie reproductive et associées à un même mâle, avec qui elles entretiennent des liens sociaux et sexuels préférentiels, et qui est souvent le protecteur et le père de leur petit. De plus, les femelles gestantes et en lactation harcèlent les femelles qui copulent avec leur mâle, probablement afin d'empêcher de nouvelles conceptions avec celui-ci. La compétition pour les soins des mâles semble donc contribuer à façonner les stratégies reproductives des femelles chez les espèces polygynes où ceux-ci apportent d'importants bénéfices aux femelles. Nous avons ensuite étudié les contraintes exercées par les mâles sur la sexualité des femelles. Mâles et femelles ont souvent des optimaux reproductifs divergents, donnant lieu à l'expression d’un conflit sexuel. Chez certaines espèces, les mâles recourent à la coercition sexuelle en agressant les femelles régulièrement afin de les obliger à s'accoupler avec eux-mêmes, ou de les empêcher de s'accoupler avec leur rivaux. Nous avons testé si l'agression dirigée par les mâles vers les femelles a une fonction de coercition sexuelle chez le babouin chacma. Nos résultats indiquent que l'agression des mâles vise en particulier les femelles sexuellement réceptives, augmente le succès d'accouplement immédiat des mâles avec la femelle harcelée et ses chances de la monopoliser lors de l'ovulation, à l'appui de l'hypothèse de coercition. Dans l'ensemble, cette étude permet d'améliorer notre compréhension des déterminants, de l'intensité, et des conséquences évolutives des contraintes sociales qui s'exercent sur la sexualité des femelles dans une société primate polygyne. Elle montre également que les conflits reproductifs jouent un rôle primordial pour structurer les relations entre les femelles d’une part, et entre les sexes d’autre part. / In group-living species, individuals of both sexes can interfere with the sexuality and social alliances of females, which may profoundly influence their reproductive strategies. Renewed attention in the operation of sexual selection on females shows that competition among females to secure reproductive resources, such as mates or allomaternal care, is common. However, to date, female reproductive competition has received little attention in polygynous species. In an attempt to fill this gap, we investigated the determinants of female reproductive competition in a polygynous primate society, the chacma baboon, focussing on a wild Namibian population. Our findings highlight that the frequency of aggression is most intense among females who are reproductively synchronous and who share the same male carer of their offspring. Females also harass sexually receptive females who attempt to mate with their offspring’s carer, likely to prevent further conceptions with him. Overall, competition to secure male carers seems to play an important role in shaping female reproductive strategies in polygynous species where males may provide females with important fitness benefits. We then examined constraints exerted by males on female sexuality. Males and females often have diverging reproductive optima, which underpins sexual conflict. In some species, males may use sexual coercion, in the form of repeated aggression before or during female sexual receptivity to induce females into mating or prevent them from mating with rivals. Here, we tested whether male aggression directed at females represents sexual coercion in chacma baboons. In support of the sexual coercion hypothesis, we found that male aggression against females is most intense when females are sexually receptive, increases male mating success with the harassed female on the short-term, and increases his chances to monopolize her around ovulation on the longer-term. Altogether, these results shed light on the determinants, intensity and evolutionary consequences of social constraints exerted on female sexuality in polygynous primates, and highlight that reproductive conflicts play a primary role in structuring female-female and male-female relationships.