This thesis explores the far-reaching impacts of sexual conflict over care on a suite of traits in the penduline tit family (Remizidae), further confirming the intricate relationships between parental care, mating systems and sexual selection. The results reveal the first genetic phylogeny of this family and suggest that uniparental care evolved once in this group. The transition to uniparental care is associated with rapid evolution of male plumage ornaments most likely driven by increased sexual selection. The results also suggest a relationship between male care and the likelihood of paternity on an evolutionary time-scale, as the biparental species exhibit much lower levels of promiscuity than the uniparental European penduline tit. Increased promiscuity was also found to impact sperm morphology in the penduline tits and allies with greater sperm length uniformity in more promiscuous species. This sperm trait was also discovered to co-vary with a sexually selected plumage trait in the European penduline tit suggesting potential interactions between female mate choice and male fertility. An investigation into genetic diversity within the Sylvioidea super-family finds large variation but does not suggest any link between promiscuity and genetic diversity as predicted if promiscuity maintains a higher effective population size in these passerines. The work highlights the interlinked relationships between parental care, mating systems and sexually selected traits, which are increasingly studied in concert. The consequences of sexual conflict over care appear to be far-reaching in the penduline tits, however the degree to which they feedback upon each other and the effect that it has on speciation remains to be seen. The penduline tits further prove their ability to provide valuable insight into the evolution of sexual conflict.

The difference in evolutionary interests of males and females can select for traits that favour an individual??s fitness at the expense of their mate??s lifetime fitness. Despite the widespread occurrence of this sexual conflict over reproductive interactions, however, research to date has largely focused on the fitness costs imposed on females by manipulative males. Empirical evidence is particularly sparse for how mating can also be costly for males, the genetic structure of traits involved in reproductive interactions, and how sexual conflict can modify sexual selection in general. My aim was to explore the broader evolutionary consequence of sexual conflict and male-female interactions. In the nuptial-feeding Australian ground cricket, Pteronemobius sp., I used an experimental evolution approach to explore how diet and sexual conflict interact to determine the costs of mating. In the Australian black field cricket, Teleogryllus commodus, I used molecular and quantitative genetic approaches to characterise the fitness consequences and genetic basis of spermatophore attachment, a trait at the centre of inter-locus sexual conflict, and then related this to both condition and male attractiveness. Finally, in T. commodus, I quantified how sexual conflict alters the sexual selection acting on male sexual traits and how this in turn shapes genetic architecture and the persistence of additive genetic variance. My results demonstrate the complex nature of reproductive interactions between males and females. Importantly, I show that diet can mediate the expression of sexual conflict in a mating system and shape the evolution of male life-span. I also show that reproductive interactions influence the fitness benefits that both male and females obtain from mating in ways that are not predicted by current theory and that much of the potential for such traits to co-evolve is via a common genetic association with condition. Finally, I demonstrate that sexual conflict can profoundly modify the process and outcome of sexual selection, thereby influencing how additive genetic variation is maintained in a suite of male sexual traits. These results highlight the need for a greater integration of sexual conflict and sexual selection theory as the evolutionary potential and significance of sexual conflict may currently be underestimated.

Sexual conflict and condition-dependent trait expression have emerged as major themes in sexual selection. There is now considerable evidence suggesting that both conflict and condition-dependence can drive the evolution and expression of sexual traits; still, important questions remain concerning the extent to which conflict shapes sexual traits and the role of condition in mediating conflict. Here, I address these two themes in studies of a ladybird mating system. One set of studies investigates the function and economics of potentially antagonistic traits – nuptial gifts and female mating resistance – while another examines condition-dependence in mating resistance and male ejaculate composition. Nuptial gifts are often considered beneficial to females, but recent thinking suggests they may also allow males to manipulate females, raising the possibility of conflict. I demonstrate that male ladybirds benefit from nuptial feeding by their mates through reduced re-mating frequency. Benefits to female reproduction and lifespan, however, are weak or non-existent. These results show that although males gain from transferring gifts that influence female behaviour, females experience neither harm nor benefit. I next tested the hypothesis that nuptial feeding is maintained – despite an absence of benefits – because female foraging is generally elevated after mating. However, although females indeed display strongly increased foraging after mating, this response did not increase nuptial feeding. Recent studies suggest that individual condition may affect the economics of mating and extent of conflict. Female ladybirds vigorously resist mating, and I show that (1) resistance is condition-dependent, with low-condition females displaying more resistance, and (2) resistance functions to minimize superfluous matings (sexual conflict), rather than to select among males (indirectly benefiting females). Resistance generates selection favouring large males; thus, this work demonstrates that ecological circumstances, through influencing condition, affect the strength of sexual selection. Finally, male condition may influence investment in ejaculate components, but condition-dependence in ejaculate composition is currently poorly understood. I show that, in agreement with theory, males in poor condition transfer smaller ejaculates that nonetheless contain more sperm, but less seminal fluid. Taken together, this work highlights both the value of economic studies in evaluating sexual conflict, and the significance of condition-dependence for sexual selection.

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.

Sexual conflict and condition-dependent trait expression have emerged as major themes in sexual selection. There is now considerable evidence suggesting that both conflict and condition-dependence can drive the evolution and expression of sexual traits; still, important questions remain concerning the extent to which conflict shapes sexual traits and the role of condition in mediating conflict. Here, I address these two themes in studies of a ladybird mating system. One set of studies investigates the function and economics of potentially antagonistic traits – nuptial gifts and female mating resistance – while another examines condition-dependence in mating resistance and male ejaculate composition. Nuptial gifts are often considered beneficial to females, but recent thinking suggests they may also allow males to manipulate females, raising the possibility of conflict. I demonstrate that male ladybirds benefit from nuptial feeding by their mates through reduced re-mating frequency. Benefits to female reproduction and lifespan, however, are weak or non-existent. These results show that although males gain from transferring gifts that influence female behaviour, females experience neither harm nor benefit. I next tested the hypothesis that nuptial feeding is maintained – despite an absence of benefits – because female foraging is generally elevated after mating. However, although females indeed display strongly increased foraging after mating, this response did not increase nuptial feeding. Recent studies suggest that individual condition may affect the economics of mating and extent of conflict. Female ladybirds vigorously resist mating, and I show that (1) resistance is condition-dependent, with low-condition females displaying more resistance, and (2) resistance functions to minimize superfluous matings (sexual conflict), rather than to select among males (indirectly benefiting females). Resistance generates selection favouring large males; thus, this work demonstrates that ecological circumstances, through influencing condition, affect the strength of sexual selection. Finally, male condition may influence investment in ejaculate components, but condition-dependence in ejaculate composition is currently poorly understood. I show that, in agreement with theory, males in poor condition transfer smaller ejaculates that nonetheless contain more sperm, but less seminal fluid. Taken together, this work highlights both the value of economic studies in evaluating sexual conflict, and the significance of condition-dependence for sexual selection.

Pre-copulatory sexual cannibalism (pre-SC), or predation of a potential mate before sperm transfer, provides an ideal model system for behavioral ecology's current focus on inter-sexual conflict. Studying the North American fishing spider (Dolomedes triton), I tested three female-benefit hypotheses for pre-SC: indirect benefits, direct benefits, and aggressive spillover. First, pre-SC may reflect a mating bias providing females with 'good-genes' benefits. By manipulating each female's options with regard to the most cited phenotypic advantage in male spiders, body size, I show that while females exhibit no bias in their attack tendency on males of different body sizes, large males mate significantly more often than small males. Second, pre-SC may be explained by direct benefits if females use it as an adaptive foraging/mating trade-off. My work provides mixed support for this idea: (i) females vary attacks according to the availability of mates, (ii) females do not vary attacks according to the availability of food, and (iii) females derive discrete fecundity benefits from consuming a male. Finally, I tested the aggressive-spillover hypothesis, which posits that pre-SC is a by-product of selection for high levels of aggression towards prey in traditional foraging contexts. Path analysis indicated intra-individual, positive correlations between aggression in foraging contexts and the mating context, thus supporting the hypothesis. I conclude by stressing that pre-SC in a given species may rarely be explained by one hypothesis, and that studies accounting for multiple benefits that fluctuate as behavioral-ecological contexts shift should give a more realistic glimpse of behavioral ecology and evolution.

Sexual selection is pervasive in nature and plays an important role in the evolution of biological diversity both within and among sexual species. However, while we have a good understanding of how competition for reproductive opportunities and mate choice can drive the evolution of exaggerated secondary sexual traits, much less is known about how sexual selection interacts with other forms of natural selection and the consequences such interactions may have for adaptation to novel environments, the purging of deleterious mutations, and population divergence/speciation. In my thesis, I carried out a series of experiments with the fruit fly Drosophila melanogaster to test hypotheses regarding the operation of sexual selection and to broaden our understanding of how sexual selection may influence adaptation and diversification. Theory suggests that natural and sexual selection may align to promote adaptation and the purging of deleterious mutations, although the harm imposed by sexual conflict may counter this. In two separate experiments, I find no evidence that sexual selection promotes adaptation to a novel environment and, rather than aligning with natural selection, I find that the effects of sexual conflict may cause sexual selection to hamper the purging of deleterious mutations. With respect to diversification, sexual conflict has been suggested to be an important, non-ecological driver of population divergence. However, the traits involved in sexual conflict may also affect nonsexual fitness and natural selection may therefore act to constrain diversification. Using an evolution experiment, I demonstrate ecologically-dependent parallel evolution of traits involved in sexual conflict, providing evidence for ecology’s importance in divergence via sexual conflict. Overall, my work has shed light on the interaction of natural and sexual selection and the consequences this may have beyond the evolution of exaggerated sexual displays and armaments.

No description available.

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.

The difference in evolutionary interests of males and females can select for traits that favour an individual??s fitness at the expense of their mate??s lifetime fitness. Despite the widespread occurrence of this sexual conflict over reproductive interactions, however, research to date has largely focused on the fitness costs imposed on females by manipulative males. Empirical evidence is particularly sparse for how mating can also be costly for males, the genetic structure of traits involved in reproductive interactions, and how sexual conflict can modify sexual selection in general. My aim was to explore the broader evolutionary consequence of sexual conflict and male-female interactions. In the nuptial-feeding Australian ground cricket, Pteronemobius sp., I used an experimental evolution approach to explore how diet and sexual conflict interact to determine the costs of mating. In the Australian black field cricket, Teleogryllus commodus, I used molecular and quantitative genetic approaches to characterise the fitness consequences and genetic basis of spermatophore attachment, a trait at the centre of inter-locus sexual conflict, and then related this to both condition and male attractiveness. Finally, in T. commodus, I quantified how sexual conflict alters the sexual selection acting on male sexual traits and how this in turn shapes genetic architecture and the persistence of additive genetic variance. My results demonstrate the complex nature of reproductive interactions between males and females. Importantly, I show that diet can mediate the expression of sexual conflict in a mating system and shape the evolution of male life-span. I also show that reproductive interactions influence the fitness benefits that both male and females obtain from mating in ways that are not predicted by current theory and that much of the potential for such traits to co-evolve is via a common genetic association with condition. Finally, I demonstrate that sexual conflict can profoundly modify the process and outcome of sexual selection, thereby influencing how additive genetic variation is maintained in a suite of male sexual traits. These results highlight the need for a greater integration of sexual conflict and sexual selection theory as the evolutionary potential and significance of sexual conflict may currently be underestimated.