Multiple mating by females is common in nature. Yet, the evolution and maintenance of polyandry remains a bit of an evolutionary puzzle. It was my aim in this thesis to reach a greater understanding of this phenomenon as well as to investigate the consequences of polyandry on the evolution of reproductive divergence in insects. In an extensive meta analysis addressing the direct effects of multiple mating on female fitness in insects, I found that insects gain from multiple matings in terms of increased lifetime offspring production. In species without nuptial feeding, increased mating rate leads to decreased female lifespan and my results strongly support the existence of an intermediate optimal female mating rate. However, results from an experimental study where I examined the relationship between female fitness and mating rate in the bean weevil (Callosobruchus maculatus) showed that female fitness was maximized at two alternative mating rates, indicating that some species may exhibit a more complex relationship between the costs and benefits of mating. In the meta analysis on species with nuptial feeding, I found only positive effects of increased mating rate and the puzzle is rather what constrains the actual mating rates of females in these groups. Sexual selection is a very potent driver of rapid evolutionary change in reproductive characters. Most research has focussed on precopulatory sexual selection, but in promiscuous species sexual selection continues after copulation and variance in male fertilization success gives rise to postcopulatory sexual selection. In this thesis I found that three allopatric populations of the red flour beetle (Tribolium castaneum) have diverged in traits related to reproduction. Male genotype affected all aspects of female reproduction, but more interestingly, males and females interacted in their effect on offspring production and reproductive rate, showing that the divergence was due to the evolution of both male and female reproductive traits. When studying postcopulatory sexual selection, sperm competition has been put forward as the main source of variance in fertilization success. The results from a set of double-mating experiments, using the same populations of flour beetles, provided strong evidence that cryptic female choice is also important in generating variance in male fertilization success. I found not only main effects of female genotype on male fertilization success but also male-female interactions which provide more unambiguous evidence for cryptic female choice. Finally, I attempted to uncover which male signals-female receptors are involved in the reproductive divergence observed in the Tribolium populations. In a double-mating experiment I manipulated female perception of two male reproductive signals, copulatory courtship and cuticular hydrocarbons, and the results indicate that, within populations, both signals are sexually selected. However, only male cuticular hydrocarbons seem to be involved in the reproductive divergence between the populations. In conclusion, multiple mating by female insects can be understood solely in terms of direct fitness benefits resulting from increased offspring production. I have shown that postcopulatory sexual selection can lead to rapid divergence in reproductive traits related to mating and that cryptic female choice plays an important role in this divergence.

Sexual conflict can lead to antagonistic coevolution between the sexes, but empirical examples are few. In this thesis secondary sexual characters in diving beetles are interpreted in the light of sexual conflict theory. Whether the male tarsal suction cups and female dorsal modifications are involved in a coevolutionary arms race is tested in two ways. First eight populations of a species with dimorphic females that varied in frequency of the morphs were investigated and male tarsal characteristics quantified. The frequency of female morphs is shown to be significantly correlated to the average number and size of male tarsal suction cups in the population, a prediction of the arms race hypothesis. Second, the hypothesis is tested in a phylogenetic perspective by optimizing the secondary sexual characters on a phylogeny. A full taxonomic revision of the genus Acilius is presented, including new synonyms, lectotype designations, geographic distributions based on more than five thousand examined museum specimens and the description of a new species from northeastern USA. Specimens of all species (except one possibly extinct that failed to be found in Yunnan, China 2000), were field collected between 2000 and 2003 in Sardinia, Sweden, Russia, Honshu and Hokkaido in Japan, New York, Maryland, California and Alberta. Three genes (CO1, H3 and Wingless) were sequenced from the fresh material as well as scoring a morphological character matrix all of which was used to derive a robust and complete hypothesis of the phylogenetic relationship in the group. The phylogeny was derived using Bayesian phylogenetics with Markov Chain Monte Carlo techniques and received a posterior probability of 0.85. Changes in male and female characters turned out to be perfectly correlated across the phylogeny, providing one of the best empirical examples to date of an antagonistic arms race between the sexes in a group of organisms. Finally, a review of a pitfall to phylogenetic analysis known under the name long-branch attraction (LBA), is provided. The problem is well known theoretically but has been questioned to occur in real data, and LBA has been in the core center of the hard debate between parsimony and likelihood advocates since different inference methods vary in sensitivity to the phenomenon. Most important conclusions from the review are; LBA is very common in real data, and is most often introduced with the inclusion of outgroups that almost always provide long branches, pulling down long terminal ingroup branches towards the root. Therefore it is recommended to always run analyses with and without outgroups. Taxon sampling is very important to avoid the pitfall as well as including different kind of data, especially morphological data, i.e. many LBA-affected conclusions have recently been reached by analyses of few taxa with complete genomes. Long-branch extraction (incl. outgroup exclusion), methodological disconcordance (parsimony vs modelbased), separate partition analyses (morphology vs molecules, codon positions, genes, etc), parametric simulation (incl. random outgroups), and split graphs are available relevant methods for the detection of LBA that should be used in combinations, because none alone is enough to stipulate LBA.

Harmony and cooperation was for long believed to dominate sexual interactions. This view slowly started to change 25 years ago and is today replaced with a view where males and females act based on what is best from a costs-benefits perspective. When sex specific costs and benefits differ, concerning reproductive decision influenced by both sexes, sexual conflict will occur. The basis for discordant reproductive interests between the sexes is that males produce many small gametes, while females’ produce few and large gametes. One result of this difference is that the optimal mating rate differs between the sexes. Males, with their many small sperm, maximize their reproductive output by mating with many females, while females often do best by not mating more frequently than to fertilize their eggs, since mating often entails a cost. Sexual conflict over mating is thus an important factor shaping the interactions between the sexes. In this thesis I study this and related conflicts between the sexes, using mathematical models, fruit flies and comparative methods. Mathematical modelling was used to explore how males and females may coevolve under sexual conflict over mating. This model shows that sexual conflict over mating results in the evolution of costly female mate choice, in terms high resistance to matings, and costly exaggerated male sexual traits, aimed to manipulate females into mating. A key assumption in this model is that males which females find attractive also are more harmful to females. This assumption was tested by housing fruit fly females with either attractive or unattractive males. Females kept with attractive males were courted and mated more, and suffered a 16 percent reduction in lifetime offspring production. In another study I measured genetic variation in two antagonistic male traits used to compete over females; offence - a male’s ability to acquire new mates and supplant stored sperm, and defence - a male’s ability to induce fidelity in his mates and prevent sperm displacement when remating occurs. Independent additive genetic variation and positive selection gradients were found for both these traits, indicating an ongoing arms race between these male antagonistic traits. This arms race also had a negative impact on females, since high values of offence compromised female fitness. Genetic variation in female ability to withstand male harm was also tested for and found, indicating that females evolve counter adaptations to reduce the effect of harmful male traits. Finally, the proposed link between sexual conflict and speciation was tested. Theory suggests that perpetual sexual arms races will cause allopatric populations to evolve along different evolutionary trajectories, resulting in speciation. This theory was tested using comparative methods by contrasting the number of extant species in taxa with high and low opportunity for sexual conflict. The study showed that taxa with high opportunity for sexual conflict, on average, has four times as many species as those with low opportunity, supporting that sexual conflict is a key process in speciation.

The ultimate goal of my thesis is to develop a better understanding of the contribution of indirect benefits (i.e. good genes) to the evolution of female mate preferences. It is genetic variance in, and genetic correlations (covariances) among, male sexual displays, female preferences for them, and fitness that in part determine the degree to which females preferring certain male displays over others will gain an indirect benefit by having higher fitness offspring. Recent advances in quantitative genetic theory provide the mathematical means for quantifying the strength of indirect selection for female mate preferences (Kirkpatrick and Hall 2004), at least under certain conditions, but there are few empirical systems for which such data exist (Brooks and Endler 2001; Qvarnström et al. 2006). I have undertaken a classic half-sibling breeding design with the ultimate goal of estimating the specific parameters of this model in a population of the Australian fruit fly Drosophila serrata. The breeding design was performed across two environments - one to which the population was well adapted and a novel environment to which it was not - thereby also providing insight into genotype-by-environment interactions for this suite of traits and their effects on good genes indirect benefits in a novel environment. General insight is also gained into the genetic covariance of male and female fitness and the prevalence of intralocus sexual conflict, the quantitative genetic basis of female mate preferences for multiple male traits, the condition-dependence of these traits, and the genetic association between sexual displays and fitness when mutation-selection balance is inferred. My results advocate caution in the application of existing theory to quantify the strength of indirect selection, suggesting that a good genes process may be fundamentally different when the exaggeration of sexual displays is eventually halted and an equilibrium is reached between opposing selection.

The ultimate goal of my thesis is to develop a better understanding of the contribution of indirect benefits (i.e. good genes) to the evolution of female mate preferences. It is genetic variance in, and genetic correlations (covariances) among, male sexual displays, female preferences for them, and fitness that in part determine the degree to which females preferring certain male displays over others will gain an indirect benefit by having higher fitness offspring. Recent advances in quantitative genetic theory provide the mathematical means for quantifying the strength of indirect selection for female mate preferences (Kirkpatrick and Hall 2004), at least under certain conditions, but there are few empirical systems for which such data exist (Brooks and Endler 2001; Qvarnström et al. 2006). I have undertaken a classic half-sibling breeding design with the ultimate goal of estimating the specific parameters of this model in a population of the Australian fruit fly Drosophila serrata. The breeding design was performed across two environments - one to which the population was well adapted and a novel environment to which it was not - thereby also providing insight into genotype-by-environment interactions for this suite of traits and their effects on good genes indirect benefits in a novel environment. General insight is also gained into the genetic covariance of male and female fitness and the prevalence of intralocus sexual conflict, the quantitative genetic basis of female mate preferences for multiple male traits, the condition-dependence of these traits, and the genetic association between sexual displays and fitness when mutation-selection balance is inferred. My results advocate caution in the application of existing theory to quantify the strength of indirect selection, suggesting that a good genes process may be fundamentally different when the exaggeration of sexual displays is eventually halted and an equilibrium is reached between opposing selection.

The ultimate goal of my thesis is to develop a better understanding of the contribution of indirect benefits (i.e. good genes) to the evolution of female mate preferences. It is genetic variance in, and genetic correlations (covariances) among, male sexual displays, female preferences for them, and fitness that in part determine the degree to which females preferring certain male displays over others will gain an indirect benefit by having higher fitness offspring. Recent advances in quantitative genetic theory provide the mathematical means for quantifying the strength of indirect selection for female mate preferences (Kirkpatrick and Hall 2004), at least under certain conditions, but there are few empirical systems for which such data exist (Brooks and Endler 2001; Qvarnström et al. 2006). I have undertaken a classic half-sibling breeding design with the ultimate goal of estimating the specific parameters of this model in a population of the Australian fruit fly Drosophila serrata. The breeding design was performed across two environments - one to which the population was well adapted and a novel environment to which it was not - thereby also providing insight into genotype-by-environment interactions for this suite of traits and their effects on good genes indirect benefits in a novel environment. General insight is also gained into the genetic covariance of male and female fitness and the prevalence of intralocus sexual conflict, the quantitative genetic basis of female mate preferences for multiple male traits, the condition-dependence of these traits, and the genetic association between sexual displays and fitness when mutation-selection balance is inferred. My results advocate caution in the application of existing theory to quantify the strength of indirect selection, suggesting that a good genes process may be fundamentally different when the exaggeration of sexual displays is eventually halted and an equilibrium is reached between opposing selection.

Les soins biparentaux impliquent que mâles et femelles travaillent ensemble et ajustent leur comportement l’un à l’autre. De nombreuses études théoriques et empiriques se sont intéressées à la façon dont un parent réagit face à une augmentation ou une diminution de l’effort parental de son partenaire. Mais très peu se sont intéressées aux mécanismes qui permettent cet ajustement. C’est chez les oiseaux qu’on trouve le plus d’espèces à soins biparentaux. Parce que ce sont des animaux qui produisent beaucoup de sons, la communication acoustique pourrait jouer un rôle dans l’organisation des soins parentaux. Le but de cette thèse est de contribuer à vérifier cette hypothèse en comparant deux espèces d’oiseaux qui diffèrent dans la façon dont le mâle et la femelle se partagent les soins parentaux. Chez le diamant mandarin, le mâle et la femelle participent tous les deux à l’incubation des oeufs tandis que c’est la femelle seule qui se charge de cette tâche chez la mésange charbonnière. Cette dernière est alors nourrie en partie par le mâle. Chez ces deux espèces, j’ai d’abord observé en milieu naturel dans quels contextes les parents communiquent avec des sons au nid etquelle est la structure de leurs échanges vocaux. J’ai pu ainsi formuler des premières hypothèses sur les fonctions de cette communication. J’ai ensuite testé expérimentalement une de ces hypothèses pour chaque espèce. Chez le diamant mandarin, j’ai montré que la communication au nid permet aux parents de se partager le temps d’incubation. Chez la mésange charbonnière la communication au nid permet à la femelle d’indiquer ses besoins en nourriture au mâle. La communication acoustique joue donc bien un rôle dans l’organisation des soins parentaux chez ces deux espèces et pourrait être un élément clef dans les études futures s’intéressant à l’ajustement de l’effort parental entre les mâles et les femelles / Biparental care involves that male and female work as a team and each individual adjusts its behaviour to that of its partner. Many theoretical and empirical studies have investigated how each parent adjusts its behaviour to an increase or a decrease in the parental effort of its partner. But fewer have explored the mechanism allowing this adjustment. Biparental care is widespread in birds. Because birds are vocal animals, acoustic communication may play a role in organizing parental care. The aim of this thesis is to contribute to answer this question by comparing two bird species that differ in sex roles during parental care. In the zebra finch, both male and female incubate the eggs, whereas in the great tit, the female incubates alone and is partly fed by the male. In both species, I first observed in which contexts parents communicate with sounds at the nest and described the structure of their vocal exchanges. This allowed me to formulate hypotheses on the functions of this communication. Then, I experimentally tested one of these hypotheses for each species. In the zebra finch, I demonstrated that communication at the nest allows the pair to share incubation time. In the great tit, communication at the nest allows the female to communicate her needs for food to her mate. Acoustic communication thus plays a role in organizing parental care in these two species and could be a key factor in future studies on the adjustment of parental effort between males and females

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.

No description available.

The ultimate goal of my thesis is to develop a better understanding of the contribution of indirect benefits (i.e. good genes) to the evolution of female mate preferences. It is genetic variance in, and genetic correlations (covariances) among, male sexual displays, female preferences for them, and fitness that in part determine the degree to which females preferring certain male displays over others will gain an indirect benefit by having higher fitness offspring. Recent advances in quantitative genetic theory provide the mathematical means for quantifying the strength of indirect selection for female mate preferences (Kirkpatrick and Hall 2004), at least under certain conditions, but there are few empirical systems for which such data exist (Brooks and Endler 2001; Qvarnström et al. 2006). I have undertaken a classic half-sibling breeding design with the ultimate goal of estimating the specific parameters of this model in a population of the Australian fruit fly Drosophila serrata. The breeding design was performed across two environments - one to which the population was well adapted and a novel environment to which it was not - thereby also providing insight into genotype-by-environment interactions for this suite of traits and their effects on good genes indirect benefits in a novel environment. General insight is also gained into the genetic covariance of male and female fitness and the prevalence of intralocus sexual conflict, the quantitative genetic basis of female mate preferences for multiple male traits, the condition-dependence of these traits, and the genetic association between sexual displays and fitness when mutation-selection balance is inferred. My results advocate caution in the application of existing theory to quantify the strength of indirect selection, suggesting that a good genes process may be fundamentally different when the exaggeration of sexual displays is eventually halted and an equilibrium is reached between opposing selection.