Populations must adapt to environmental changes to remain viable. Both evolution and phenotypic plasticity contribute to adaptation, with plasticity possibly being more important for coping with rapid change. Adaptation is complex in species with separate sexes, as the sexes can differ in the strength or direction of natural selection, the genetic basis of trait variation, and phenotypic plasticity. Many species show sex differences in plasticity, yet how these differences influence extinction susceptibility remains unclear. We first extend theoretical models of population persistence in changing environments and show that persistence is affected by sexual dimorphism for phenotypic plasticity, trait genetic architecture, and sex-specific selection. Our models predict that female-biased adaptive plasticity-particularly in traits with modest-to-low cross-sex genetic correlations-typically promotes persistence, though we also identify conditions where sexually monomorphic or male-biased plasticity promotes persistence. We then perform a meta-analysis of sex-specific plasticity under manipulated thermal conditions. Although examples of sexually dimorphic plasticity are widely observed, systematic sex differences are rare. An exception-cold resistance-is systematically female-biased and represents a trait wherein sexually dimorphic plasticity might elevate population viability in changing environments. We discuss our results in light of debates about the roles of evolution and plasticity in extinction susceptibility.

Facultative sexual organisms must allocate resources to both asexual and sexual reproduction. Optimal patterns of investment in sex depend on the relative costs and benefits of each reproductive mode, and may consequently be context- and condition-dependent. Two proposed explanations for the observed variation in investment in sex among facultative sexual lineages invoke alternative condition-dependent scenarios. Under the fitness-associated sex hypothesis, sex is predicted when individuals are in poor condition or experience stressful environments. Under the resource-demanding sex hypothesis, sex is only affordable to individuals in good condition experiencing favourable environments. Direct tests of these contrasting hypotheses are rare; moreover, investment in different components of sexual reproduction responds differently to cues promoting sex, and may be subject to different energetic constraints. Using genotypes of facultative sexual Daphnia carinata that differ in their level of investment in sex, we manipulated resource availability while accounting for day length (a seasonal cue for sex) to evaluate these hypotheses. The sexual response to day length depended on resource availability: increased day lengths and reduced food availability increased the production of sexual eggs, and relative investment in males, in a manner consistent with the fitness-associated sex hypothesis. The pattern of condition-dependence was specific to each component of reproductive investment – while male production covaried with asexual fecundity across genotypes, increased sexual egg production was associated with reduced asexual reproduction. Our results suggest that investment in sex is determined largely by its context-dependent advantages, and that this investment is not moderated by immediate costs to asexual reproduction.

Research predicting the impact and spread of infectious disease has been heavily influenced by the idea of an evolutionary trade-off between a pathogen's virulence and its transmission rate. In a meta-analysis of the key underlying relationships, Acevedo etal. (2019) highlight the surprising lack of empirical evidence for this influential hypothesis.

Adaptive explanations for dormancy often invoke bet hedging, where reduced mean fitness can be adaptive if it associates with reduced fitness variance. Sex allocation theory typically ignores variance effects and focuses on mean fitness. For many cyclical parthenogens, these themes become linked, as only sexually produced eggs undergo the dormancy needed to survive harsh conditions. We ask how sex allocation and the timing of sex evolve when this constraint exists in the form of a trade-off between asexual reproduction and sexual production of dormant eggs-the former being crucial for within-season success and the latter for survival across seasons. We show that male production can be temporally separated from or co-occur with sex, depending on whether direct (time) or indirect (population density) cues of the season's end are available and whether population growth is density dependent. Sex generally occurs late in the season but is induced earlier in unpredictable environments. When only indirect cues are available, the temporal spread of sex, and with it the production of dormant stages, is even larger and, given sufficient mortality, leads to endogenous population cycles in which frequent sex coincides with high densities. In all scenarios, algorithms maximizing geometric mean fitness have reduced fitness variance compared with a hypothetical non-bet hedger, confirming that the timing of male production and sex in facultative seasonal settings can be bet-hedging traits.

Local adaptation is of fundamental interest to evolutionary biologists. Traditionally, local adaptation has been studied using reciprocal transplant experiments to quantify fitness differences between residents and immigrants in pairwise transplants between study populations. Previous studies have detected local adaptation in some cases, but others have shown lack of adaptation or even maladaptation. Recently, the importance of different fitness components, such as survival and fecundity, to local adaptation have been emphasized. Here, we address another neglected aspect in studies of local adaptation: sex differences. Given the ubiquity of sexual dimorphism in life histories and phenotypic traits, this neglect is surprising, but may be partly explained by differences in research traditions and terminology in the fields of local adaptation and sexual selection. Studies that investigate differences in mating success between resident and immigrants across populations tend to be framed in terms of reproductive and behavioural isolation, rather than local adaptation. We briefly review the published literature that bridges these areas and suggest that reciprocal transplant experiments could benefit from quantifying both male and female fitness components. Such a more integrative research approach could clarify the role of sex differences in the evolution of local adaptations.This article is part of the theme issue 'Linking local adaptation with the evolution of sex differences'.

The timing of sex in facultatively sexual organisms is critical to fitness, due to the differing demographic consequences of sexual versus asexual reproduction. In addition to the costs of sex itself, an association of sex with the production of dormant life stages also influences the optimal use of sex, especially in environments where resting eggs are essential to survive unfavourable conditions. Here we document population dynamics and the occurrence of sexual reproduction in natural populations of Daphnia magna across their growing season. The frequency of sexually reproducing females and males increased with population density and with decreasing asexual clutch sizes. The frequency of sexually reproducing females additionally increased as population growth rates decreased. Consistent with population dynamic models showing that the opportunity cost of sexual reproduction (foregoing contribution to current population growth) diminishes as populations approach carrying capacity, we found that investment in sexual reproduction was highest when asexual population growth was low or negative. Our results support the idea that the timing of sex is linked with periods when the relative cost of sex is reduced due to low potential asexual growth at high population densities. Thus, a combination of ecological and demographic factors affect the optimal timing of sexual reproduction, allowing D. magna to balance the necessity of sex against its costs.

Cyclical parthenogenesis presents an interesting challenge for the study of sex allocation, as individuals' allocation decisions involve both the choice between sexual and asexual reproduction, and the choice between sons and daughters. Male production is therefore expected to depend on ecological and evolutionary drivers of overall investment in sex, and those influencing male reproductive value during sexual periods. We manipulated experimental populations, and made repeated observations of natural populations over their growing season, to disentangle effects of population density and the timing of sex from effects of adult sex ratio on sex allocation in cyclically parthenogenetic Daphnia magna. Male production increased with population density, the major ecological driver of sexual reproduction; however, this response was dampened when the population sex ratio was more male-biased. Thus, in line with sex ratio theory, we show that D. magna adjust offspring sex allocation in response to the current population sex ratio.

Sex allocation theory predicts that females mated to attractive males produce more sons than females mated to unattractive males. However, previous tests of this hypothesis have obtained mixed results. We suggest that females differ in the traits they find attractive. To test this proposition, we assessed female zebra finches' preferences for males banded with red or green plastic leg bands and then tested the sex allocation pattern of females paired with preferred and non-preferred males. Although most females preferred red-banded males, 34% of females consistently preferred green-banded males. The sex ratio at laying and hatching was not influenced by paternal preferred status. However, the fledging sex ratios differed between females paired with preferred and non-preferred males due to sex-biased chick mortality; sons of females paired with preferred males were born heavier and were more likely to survive than daughters. Our results indicate that female zebra finches show individual variation in their preferences for colour banded males, although females do not seem to adjust the offspring sex ratio in response to their mate's band colour. However, the differential post-hatch mortality suggests females may differentially allocate resources into male and female eggs according to their individual mate preferences.

Sexual selection imposed by mating preferences is often implicated in the evolution of both sexual dimorphism and divergence between species in signalling traits. Epicuticular compounds (ECs) are important signalling traits in insects and show extensive variability among and within taxa. Here, we investigate whether variation in the multivariate EC profiles of two sex role‐reversed beetle species, Megabruchidius dorsalis and Megabruchidius tonkineus, predicts mate attractiveness and mating success in males and females. The two species had highly distinct EC profiles and both showed significant sexual dimorphism in ECs. Age and mating status in both species were also distinguishable by EC profile. Males and females of both species showed significant association between their EC profile and attractiveness, measured both as latency to mating and as success in mate‐choice trials. Remarkably, the major multivariate vector describing attractiveness was correlated in both species, both sexes, and in both choice and no‐choice experiments such that increased attractiveness was in all cases associated with a similar multivariate modification of EC composition. Furthermore, in both sexes this vector of attractiveness was associated with more male‐like EC profiles, as well as those characterizing younger and nonvirgin individuals, which might reflect a general preference for individuals of high condition in both sexes. Despite significant sexual selection on EC composition, however, we found no support for the proposition that sexual selection is responsible for divergence in ECs between these species.

The reversal of conventional sex roles was enigmatic to Darwin, who suggested that it may evolve when sex ratios are female biased [1]. Here we present direct evidence confirming Darwin's hypothesis. We investigated mating system evolution in a sex-role-reversed beetle (Megabruchidius dorsalis) using experimental evolution under manipulated sex ratios and food regimes. In female-biased populations, where reproductive competition among females was intensified, females evolved to be more attractive and the sex roles became more reversed. Interestingly, female-specific mating behavior evolved more rapidly than male-specific mating behavior. We show that sexual selection due to reproductive competition can be strong in females and can target much the same traits as in males of species with conventional mating systems. Our study highlights two central points: the role of ecology in directing sexual selection and the role that females play in mating system evolution.

Male genitalia often show remarkable differences among related species in size, shape and complexity. Across poeciliid fishes, the elongated fin (gonopodium) that males use to inseminate females ranges from 18 to 53% of body length. Relative genital size therefore varies greatly among species. In contrast, there is often tight within-species allometric scaling, which suggests strong selection against genital–body size combinations that deviate from a species’ natural line of allometry. We tested this constraint by artificially selecting on the allometric intercept, creating lines of males with relatively longer or shorter gonopodia than occur naturally for a given body size in mosquitofish, Gambusia holbrooki . We show that relative genital length is heritable and diverged 7.6–8.9% between our up-selected and down-selected lines, with correlated changes in body shape. However, deviation from the natural line of allometry does not affect male success in assays of attractiveness, swimming performance and, crucially, reproductive success (paternity).

In many birds, males are presumed to protect their paternity by closely guarding their mate or copulating frequently with her. Both these costly behaviors are assumed to reduce the risk and/or intensity of sperm competition. However, despite many studies on avian extra-pair paternity, it remains unclear how strongly these behaviors are related to fitness and other key life-history traits. Here, we conduct meta-analyses to address two questions. First, are mate guarding and/or frequent copulation positively correlated with a male's share of paternity at his nest? We find a significant positive correlation between both presumed paternity protection behaviors and paternity share. The relationship is, however, weak (r = 0.08-0.23). This is perhaps unsurprising if the risk of partner infidelity, hence the need to protect paternity, varies among males. For example, more attractive males might have less need to protect their paternity. Second, do males with higher indices of so-called male "quality" (phenotypic measures, usually subjectively defined by researchers as predictors of male attractiveness) exhibit lower levels of paternity protection behavior? We find a negative correlation between male quality and paternity protection. This finding might partly explain the weak relationship between paternity protection and paternity, although we discuss other, nonmutually exclusive possibilities.

The operational sex ratio (OSR: sexually active males: receptive females) predicts the intensity of competition for mates. It is less clear, however, under what circumstances, the OSR predicts the strength of sexual selection - that is, the extent to which variation in mating success is attributable to traits that increase the bearer's attractiveness and/or fighting ability. To establish causality, experiments that manipulate the OSR are required. Furthermore, if it is possible to control for any OSR-dependent changes in the chosen sex (e.g. changes in male courtship), we can directly test whether the OSR affects the behaviour of the choosing sex (e.g. female choice decisions). We conducted female mate choice experiments in the field using robotic models of male fiddler crabs (Uca mjoebergi). We used a novel design with two females tested sequentially per trial. As in nature, the choice of the first female to mate therefore affected the mates available to the next female. In general, we detected significant sexual selection due to female choice for 'males' with larger claws. Importantly, the strength of sexual selection did not vary across five different OSR/density treatments. However, as the OSR decreased (hence the number of available males declined), females chose the 'males' with the largest claws available significantly more often than expected by chance. Possible reasons for this mismatch between the expected and observed effects of the OSR on the strength of sexual selection are discussed.

Male harassment is a classic example of how sexual conflict over mating leads to sex-specific behavioural adaptations. Females often suffer significant costs from males attempting forced copulations, and the sexes can be in an arms race over male coercion. Yet, despite recent recognition that divergent sex-specific interests in reproduction can affect brain evolution, sexual conflict has not been addressed in this context. Here, we investigate whether artificial selection on a correlate of male success at coercion, genital length, affects brain anatomy in males and females. We analysed the brains of eastern mosquitofish (Gambusia holbrooki), which had been artificially selected for long or short gonopodium, thereby mimicking selection arising from differing levels of male harassment. By analogy to how prey species often have relatively larger brains than their predators, we found that female, but not male, brain size was greater following selection for a longer gonopodium. Brain subregion volumes remained unchanged. These results suggest that there is a positive genetic correlation between male gonopodium length and female brain size, which is possibly linked to increased female cognitive ability to avoid male coercion. We propose that sexual conflict is an important factor in the evolution of brain anatomy and cognitive ability.

Sexual selection is the prime evolutionary force that makes males and females different. This process has long been viewed as one where male compete with one another and where females choose. However, since the discovery that multiple mating by females is common in animals, sexual selection theory has been expanded to include mate competition between females and mate choice by males. However, empirical studies addressing these themes are scarce. In my thesis, I explore the evolution of sex role reversed mating systems using the honey locust beetles (Megabruchidius dorsalis and M. tonkineus). I used these species to shed light on (1) how closely sexual selection in females resembles its better‑studied male counterpart, (2) the implications of male mating costs for mating system evolution and (3) the effects of reproductive competition between females on the evolution of female courtship behaviour. By manipulating male mating rate, I found that males that mated more lived shorter lives, showing that mating is costly for males. I also demonstrated that males are choosy about whom they mate with and prefer vigorously courting females (Paper II). In contrast to males, previous studies suggested that female honey locust beetles benefit nutritionally from mating due to the large ejaculates provided by males. I manipulated male condition to show that male adult feeding had significant effects on female reproduction. Females that mated with males of good condition lived longer and produced more offspring than females whose mates were in poor condition (Paper III). When mating is costly for males, theory predicts that sexual selection in females can be strong. I compared sexual selection in honey locust beetles to that in two other species of seed beetles with conventional sex roles. I found substantial sexual selection in honey locust beetle females, which was comparable in strength to that in males (Paper I). I also measured the evolutionary effects of altered sex ratios on mating system parameters in both honey locust beetle species, using an experimental evolution design. Under female-biased sex ratios, representing strong sexual selection in females, females of M. dorsalis rapidly evolved elevated courtship intensity, thereby intensifying the reversal of sex roles (Paper V). In M. tonkineus, males evolved under male-biased sex ratios to transfer larger ejaculates, demonstrating the role of male-male reproductive competition for the evolution of male provisioning (Paper IV). My thesis highlights the essential, and often overlooked, role that females play in mating system evolution and that their contribution cannot simply be reduced to mate choice.

Females can benefit from mate choice for male traits (e.g. sexual ornaments or body condition) that reliably signal the effect that mating will have on mean offspring fitness. These male-derived benefits can be due to material and/or genetic effects. The latter include an increase in the attractiveness, hence likely mating success, of sons. Females can potentially enhance any sex-biased benefits of mating with certain males by adjusting the offspring sex ratio depending on their mate's phenotype. One hypothesis is that females should produce mainly sons when mating with more attractive or higher quality males. Here we perform a meta-analysis of the empirical literature that has accumulated to test this hypothesis. The mean effect size was small (r = 0.064-0.095; i.e. explaining <1% of variation in offspring sex ratios) but statistically significant in the predicted direction. It was, however, not robust to correction for an apparent publication bias towards significantly positive results. We also examined the strength of the relationship using different indices of male attractiveness/quality that have been invoked by researchers (ornaments, behavioural displays, female preference scores, body condition, male age, body size, and whether a male is a within-pair or extra-pair mate). Only ornamentation and body size significantly predicted the proportion of sons produced. We obtained similar results regardless of whether we ran a standard random-effects meta-analysis, or a multi-level, Bayesian model that included a correction for phylogenetic non-independence. A moderate proportion of the variance in effect sizes (51.6-56.2%) was due to variation that was not attributable to sampling error (i.e. sample size). Much of this non-sampling error variance was not attributable to phylogenetic effects or high repeatability of effect sizes among species. It was approximately equally attributable to differences (occurring for unknown reasons) in effect sizes among and within studies (25.3, 22.9% of the total variance). There were no significant effects of year of publication or two aspects of study design (experimental/observational or field/laboratory) on reported effect sizes. We discuss various practical reasons and theoretical arguments as to why small effect sizes should be expected, and why there might be relatively high variation among studies. Currently, there are no species where replicated, experimental studies show that mothers adjust the offspring sex ratio in response to a generally preferred male phenotype. Ultimately, we need more experimental studies that test directly whether females produce more sons when mated to relatively more attractive males, and that provide the requisite evidence that their sons have higher mean fitness than their daughters.

When males provide females with resources at mating, they can become the limiting sex in reproduction, in extreme cases leading to the reversal of typical courtship roles. The evolution of male provisioning is thought to be driven by male reproductive competition and selection for female fecundity enhancement. We used experimental evolution under male- or female-biased sex ratios and limited or unlimited food regimes to investigate the relative roles of these routes to male provisioning in a sex role-reversed beetle, Megabruchidius tonkineus, where males provide females with nutritious ejaculates. Males evolving under male-biased sex ratios transferred larger ejaculates than did males from female-biased populations, demonstrating a sizeable role for reproductive competition in the evolution of male provisioning. Although larger ejaculates elevated female lifetime offspring production, we found little evidence of selection for larger ejaculates via fecundity enhancement: males evolving under resource-limited and unlimited conditions did not differ in mean ejaculate size. Resource limitation did, however, affect the evolution of conditional ejaculate allocation. Our results suggest that the resource provisioning that underpins sex role reversal in this system is the result of male-male reproductive competition rather than of direct selection for males to enhance female fecundity.