Limits to Nuptial Gift Production by Male Fireflies, Photinus ignitus

Males of diverse insect species provide females with nuptial gifts, and limits on males' ability to produce these gifts may influence courtship behavior and mating systems. In the firefly Photinus ignitus, males transfer a complex spermatophore to females during mating. We provided firefly males unlimited access to responsive females to examine whether spermatophore production limits male mating success. Male spermatophore mass decreased significantly across sequential matings, and the percentage of successful matings declined during the second half of each male's life span. Male body mass explained a significant proportion of variation in size of the first spermatophore produced by P. ignitus males, but this relationship disappeared with second spermatophores. This study indicates that males' ability to produce spermatophores declines over their lifetime and that limits on nuptial giftproduction can constrain male mating success in Photinus fireflies.     

Male mating costs in a polygynous mosquito with ornaments expressed in both sexes

Male mate choice in species with conventional sex roles is difficult to explain and has, therefore, been the focus of many recent theoretical models. These models have focused on variance in female quality and, to a lesser extent, male investments/costs associated with mating. In this study, we investigate the costs of courtship and copulation in the polygynous mosquito Sabethes cyaneus. In this species, both males and females possess elaborate ornaments. Previous studies suggest that the most likely explanation for the presence of these ornaments is mutual mate choice. Thus, this system provides an excellent model for exploring the evolution of mutual mate choice in polygynous species. We disentangle the costs of courtship and copulation by monitoring male survival in three groups of males: housed alone (group 1); able only to court females (group 2); or able to court and copulate with females (group 3). We show that males incur a cost of courtship and copulation and that courtship intensity is negatively related to male longevity. Our results suggest that courtship and copulation carry additive costs to males. We discuss the implications of these results in the context of current mutual mate choice theory and suggest that courtship costs may be an unappreciated key factor in the evolution of male mate choice.     

Parental investment, sexual selection and sex ratios

Conventional sex roles imply caring females and competitive males. The evolution of sex role divergence is widely attributed to anisogamy initiating a self‐reinforcing process. The initial asymmetry in pre‐mating parental investment (eggs vs. sperm) is assumed to promote even greater divergence in post‐mating parental investment (parental care). But do we really understand the process? Trivers [Sexual Selection and the Descent of Man 1871–1971 (1972), Aldine Press, Chicago] introduced two arguments with a female and male perspective on whether to care for offspring that try to link pre‐mating and post‐mating investment. Here we review their merits and subsequent theoretical developments. The first argument is that females are more committed than males to providing care because they stand to lose a greater initial investment. This, however, commits the ‘Concorde Fallacy’ as optimal decisions should depend on future pay‐offs not past costs. Although the argument can be rephrased in terms of residual reproductive value when past investment affects future pay‐offs, it remains weak. The factors likely to change future pay‐offs seem to work against females providing more care than males. The second argument takes the reasonable premise that anisogamy produces a male‐biased operational sex ratio (OSR) leading to males competing for mates. Male care is then predicted to be less likely to evolve as it consumes resources that could otherwise be used to increase competitiveness. However, given each offspring has precisely two genetic parents (the Fisher condition), a biased OSR generates frequency‐dependent selection, analogous to Fisherian sex ratio selection, that favours increased parental investment by whichever sex faces more intense competition. Sex role divergence is therefore still an evolutionary conundrum. Here we review some possible solutions. Factors that promote conventional sex roles are sexual selection on males (but non‐random variance in male mating success must be high to override the Fisher condition), loss of paternity because of female multiple mating or group spawning and patterns of mortality that generate female‐biased adult sex ratios (ASR). We present an integrative model that shows how these factors interact to generate sex roles. We emphasize the need to distinguish between the ASR and the operational sex ratio (OSR). If mortality is higher when caring than competing this diminishes the likelihood of sex role divergence because this strongly limits the mating success of the earlier deserting sex. We illustrate this in a model where a change in relative mortality rates while caring and competing generates a shift from a mammalian type breeding system (female‐only care, male‐biased OSR and female‐biased ASR) to an avian type system (biparental care and a male‐biased OSR and ASR).     

Sexual Selection and Mate Choice

After a long period of dormancy, Darwin's theory of sexual selection in general, and mate choice in particular, now represents one of the most active fields in evolutionary research. After a brief overview of the history of ideas and a short introduction into the main mechanisms of sexual selection, I discuss some recent theoretical developments and empirical findings in the study of mate choice and review the various current models of mate choice, which can be grossly divided into adaptive models and nonadaptive models. I also examine whether available primate evidence supports various hypotheses concerning mate choice. Although primatologists were long aware that nonhuman primates have preferences for certain mating partners, until recently the functions and evolutionary consequences of their preferences remained obscure. Now there is growing evidence that mate choice decisions provide primates with important direct or indirect benefits. For example, several observations are consistent with the hypothesis that by direct or indirect mate choice female primates lower the risk of infanticide or enhance the chance of producing viable offspring. Nevertheless, there are also significant holes in our knowledge. How the male mandrill, one of Darwin's famous examples, got his brightly colored face, is still unknown.     

Sexual dimorphism of body size and shell shape in European tortoises

Adult body size and shape were examined in almost 1400 individuals of the tortoises Testudo graeca , T. hermanni and T. marginata from Greece. The size at maturity was greater in females than in males in all three species. Maximum and mean adult sizes were also greater in females than in males in T. graeca and T. hermanni . Males grew to a larger size than females in T. marginata , and mean adult size was similar in the sexes in this species. Sexual dimorphism of shape (adjusted for size covariate) was shown in most of the characters examined, and the degree of this dimorphism differed significantly among the three species. Differences were related to their contrasting courtship behaviours: horizontal head movements and severe biting in T. marginata , vertical head bobs and carapace butting in T. graeca , and mounting and tail thrusting in T. hermanni . There was no difference in the frequency of observations of courtship or fighting among the three species, but courtship was about 10 times more common than combat in males. All species showed greatest courtship activity in autumn; copulation was rarely observed in T. hermanni (only 0.36% of courting males) and not seen in the other species in the field. Observations made throughout the activity season indicated that feeding was equally common in males and females in all three species. Differences in shape were more likely to be the result of sexual selection than of natural selection for fecundity. Detailed predictions are made for sexual dimorphism of other characters in these species.     

Limits to runaway sexual selection: The wallflower paradox

In his mathematical treatment of Fisher's ideas on sexual selection (so-called runaway selection) Lande (1981) predicted that males may evolve increasingly elaborate sexual characters despite opposing viability selection as a consequence of the associated costs. Lande thereby assumed that female mate preferences are not subject to selection since (1) females are all inseminated and (2) the quantity and quality of their offspring are independent of the female's mate preferences. Kirkpatrick (1985) removed the latter assumption and investigated the consequences for the mean phenotype with respect to both female and male traits. He also explored the dynamics of the (co)-variance matrix by numerical methods. In this paper we consider a simpler model with just two multi-allelic loci. This enables us to derive explicit expressions for (co)-variances under steady state conditions. Rather than assume natural selection through differential fertility (as in Kirkpatrick, 1985), we take sexual selection on females into account by modelling the preference-dependent risk that females remain unmated. We argue that this wallflower effect is a realistic feature of any mating system, since it merely depends on the existence of (1) variation in mating preferences and (2) a finite mating season. Our approach provided an insight into the dynamic behaviour of the means of the phenotypes. This is because the dynamics of the means depend on the steady state (co)-variance matrix. Thus, an insight into the former requires explicit expressions for the latter. Whereas Lande and Kirkpatrick predicted runaway processes, despite opposing viability selection, our model predicts a globally stable steady state, i.e. no runaway, even without opposing viability selection (under the assumption of an asymptotically stable steady state of the (co)-variances. Admittedly, we have no analytic proof of this stability but only support for it, based on simulations.) The absence of the runaway processes in our model is caused by the wallflower effect, since it imposes constraints on the steady state of the (co)-variance matrix. When mutational input applies to female traits but not to male traits, explicit expressions for the (co)-variances under steady state conditions can be derived, and these show that: (1) both the genetic covariance and the variance of male traits are equal to zero, but (2) the variance of the female trait exceeds to zero. Should there be mutational input influencing the male trait, then these results would suggest that the male-to-female ratio of variances is much smaller than unity. This prediction is of tremendous importance for speciation through founding events.