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Understanding the pleiotropic consequences of gene drive systems on host fitness is essential to predict their spread through a host population. Here, we study sex-ratio (SR) X-chromosome drive in the fly Drosophila recens, where SR causes the death of Y-bearing sperm in male carriers. SR males only sire daughters, which all carry SR, thus giving the chromosome a transmission advantage. The prevalence of the SR chromosome appears stable, suggesting pleiotropic costs. It was previously shown that females homozygous for SR are sterile, and here, we test for additional fitness costs of SR.We found that females heterozygous for SR have reduced fecundity and that male SR carriers have reduced fertility in conditions of sperm competition. We then use our fitness estimates to parametrize theoretical models of SR drive and show that the decrease in fecundity and sperm competition performance can account for the observed prevalence of SR in natural populations. In addition, we found that the expected equilibrium frequency of the SR chromosome is particularly sensitive to the degree of multiple mating and performance in sperm competition. Together, our data suggest that the mating system of the organism should be carefully considered during the development of gene drive systems.

Joint nesting by females and cooperative polyandry—cooperatively breeding groups with a male-biased breeder sex ratio—are little-understood, rare breeding systems. We tested alternative hypotheses of factors potentially driving these phenomena in a population of joint-nesting acorn woodpeckers (Melanerpes formicivorus). During periods of high population density and thus low independent breeding opportunities, acorn woodpecker females formed joint-nesting coalitions with close kin. Coalitions were typically associated with groups with a male bias. We found strong evidence for both inter- and intrasexual conflict, as joint nesting conferred a fitness benefit to some males, a significant fitness cost to females, and no gain in per capita reproductive output for either sex. Such conflict, particularly the cost to females, may be an important reason why joint nesting is rare among cooperatively breeding taxa.

Broadcast spawning invertebrates offer highly tractable models for evaluating sperm competition, gamete-level mate choice and sexual conflict. By displaying the ancestral mating strategy of external fertilization, where sexual selection is constrained to act after gamete release, broadcast spawners also offer potential evolutionary insights into the cascade of events that led to sexual reproduction in more ‘derived’ groups (including humans). Moreover, the dynamic reproductive conditions faced by these animals mean that the strength and direction of sexual selection on both males and females can vary considerably. These attributes make broadcast spawning invertebrate systems uniquely suited to testing, extending, and sometimes challenging classic and contemporary ideas in sperm competition, many of which were first captured in Parker's seminal papers on the topic. Here, we provide a synthesis outlining progress in these fields, and highlight the burgeoning potential for broadcast spawners to provide both evolutionary and mechanistic understanding into gamete-level sexual selection more broadly across the animal kingdom. This article is part of the theme issue ‘Fifty years of sperm competition’.

Over the last decades, the field of sexual selection underwent a paradigm shift from sexual-stereotype thinking of \"eager\" males and \"coy\" females towards a more nuanced perspective acknowledging that not only males but also females can benefit from multiple mating and compete for mating partners. Yet, sexual selection in females is still considered a peculiarity, and the evolution of polyandry is often viewed to result from a higher mating interest of males. Here, we present meta-analytic evidence from 77 species across a broad range of animal taxa to demonstrate that female reproductive success is overall positively correlated with mating success, suggesting that females typically benefit from multiple mating. Importantly, we found that these fitness gains likely promote the evolution of polyandry. Our findings offer support for the idea that sexual selection is widespread in females and to play a key role for the evolution of animal mating systems. Thereby, our results extend our understanding of the evolutionary consequences of sexual reproduction and contribute to a more balanced view of how sexual selection operates in males and females.

Selfish genetic elements (SGEs) are diverse and near ubiquitous in Eukaryotes and can be potent drivers of evolution. Here, we discuss SGEs that specifically act on sperm to gain a transmission advantage to the next generation. The diverse SGEs that affect sperm often impose costs on carrier males, including damaging ejaculates, skewing offspring sex ratios and in particular reducing sperm-competitive success of SGE-carrying males. How males and females tolerate and mitigate against these costs is a dynamic and expanding area of research. The intense intra-genomic conflict that these selfish elements generate could also have implications for male fertility and spermatogenesis more widely. This article is part of the theme issue ‘Fifty years of sperm competition’.

We give a historic overview and critical perspective of polyandry in the context of sexual selection. Early approaches tended to obfuscate the fact that the total matings (copulations) by the two sexes is equal, neglecting female interests and that females often mate with (or receive ejaculates from) more than one male (polyandry). In recent years, we have gained much more insight into adaptive reasons for polyandry, particularly from the female perspective. However, costs and benefits of multiple mating are unlikely to be equal for males and females. These must be assessed for each partner at each potential mating between male i and female j, and will often be highly asymmetric. Interests of i and j may be in conflict, with (typically, ultimately because of primordial sex differences) i benefitting and j losing from mating, although theoretically the reverse can also obtain. Polyandry reduces the sex difference in Bateman gradients, and the probability of sexual conflict over mating by: (i) reducing the potential expected value of each mating to males in inverse proportion to the number of mates per female per clutch, and also often by (ii) increasing ejaculate costs through increased sperm allocation. It can nevertheless create conflict over fertilization and increase conflict over parental investment. The observed mean mating frequency for the population (and hence the degree of polyandry) is likely, at least in part, to reflect a resolution of sexual conflict. Immense diversity exists across and within taxa in the extent of polyandry, and views on its significance have changed radically, as we illustrate using avian polyandry as a case study. Despite recent criticisms, the contribution of the early pioneers of sexual selection, Darwin and Bateman, remains generally valid, and should not, therefore, be negated; as with much in science, pioneering advances are more often amplified and refined, rather than replaced with entirely new paradigms.

Offspring and breeding (operational) sex ratios (OSR) are a key component of demographic studies. While offspring sex ratios are often relatively easy to measure, measuring OSRs is often far more problematic. Yet, highly skewed OSRs, and a lack of male–female encounters, may be an important extinction driver. Using loggerhead sea turtles (Caretta caretta) as a case study, we showed the utility of drones, i.e. unmanned aerial vehicles (UAVs), to distinguish adult males and females in a marine breeding area, using a combination of morphological characteristics (tail length) and behavioural differences (active mating, courting and searching by males versus resting by females). Through repeated surveys, we documented seasonal changes in the OSR. While the number, and ratio, of males and females on the breeding grounds changed massively, the ratio of receptive females (derived from the rate of influx of new individuals to the area) to breeding males remained close to 1:1 for much of the period before nesting commenced. Hence, we show how large imbalances in the number of adult males and females may translate into relatively balanced OSRs. Our results suggest that the departure of males from the breeding grounds is linked to a decline in female receptivity, with female sea turtles being known to store sperm to ensure high clutch fertility throughout the nesting season. In conclusion, while we detected up to three times more females than males at the breeding ground, at present, OSRs appear stable. However, because most males breed annually (vs. biannually by females), there might only be c. 100 males in the adult population (i.e. adult sex ratio of 1:7.5), which might become further skewed under expected climate change scenarios; thus, we need to identify the minimum number of males required to prevent extinction. Finally, we highlight the use of UAVs for assessing the mating dynamics of other marine, terrestrial or avian species, in which adults might exhibit visually detectable differences, such as sexual dimorphism, external body characteristics or grouping tendencies. A plain language summary is available for this article. Plain Language Summary

Despite its potential for providing a deeper understanding of both evolutionary processes and epidemic dynamics, the reciprocal interrelationship between mating system evolution and sexually transmitted diseases remains largely unexplored. Here we developed an agent-based model simulating the evolution of two different female mating strategies (monandry versus polyandry) under the spread of a hypothetical sexually transmitted disease implying reproductive costs. Our results strongly support the existence of feedback loops between the pathogen’s transmissibility and the evolution of mating strategies. Importantly, we found several unexpected, non-linear emerging behaviours of the system, as well as tipping points which were undocumented under the conditions we considered here: i) medium-high/high probabilities of disease transmission per sexual contact reveal switches between disease-free and endemic outcomes, ii) counterintuitively, the disappearance of the pathogen might be a good indicator and predictor of the imminent extinction of the polyandrous genotype/phenotype/strategy from the population, and iii) probabilities of transmission above a medium-low threshold can offset the spread of polyandry even when this behaviour entails pronounced benefits. More broadly, our results illustrate that research into reciprocal influences between the dynamics of disease spread and sexual behaviour can provide valuable insights into disease transmission and the evolution of reproductive strategies, as well as into the sensitivity of mating systems’ evolution to small variations in ecological and disease contexts.