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Males and females often have different roles in reproduction, although the origin of these differences has remained controversial. Explaining the enigmatic reversed sex roles where males sacrifice their mating potential and provide full parental care is a particularly long-standing challenge in evolutionary biology. While most studies focused on ecological factors as the drivers of sex roles, recent research highlights the significance of social factors such as the adult sex ratio. To disentangle these propositions, here, we investigate the additive and interactive effects of several ecological and social factors on sex role variation using shorebirds (sandpipers, plovers, and allies) as model organisms that provide the full spectrum of sex role variation including some of the best-known examples of sex-role reversal. Our results consistently show that social factors play a prominent role in driving sex roles. Importantly, we show that reversed sex roles are associated with both male-skewed adult sex ratios and high breeding densities. Furthermore, phylogenetic path analyses provide general support for sex ratios driving sex role variations rather than being a consequence of sex roles. Together, these important results open future research directions by showing that different mating opportunities of males and females play a major role in generating the evolutionary diversity of sex roles, mating system, and parental care. [ABSTRACT FROM AUTHOR]

Sexual dimorphism can evolve in response to sex-specific selection pressures that vary across habitats. We studied sexual differences in subterranean amphipods Niphargus living in shallow subterranean habitats (close to the surface), cave streams (intermediate), and cave lakes (deepest and most isolated). These three habitats differ because at greater depths there is lower food availability, reduced predation, and weaker seasonality. Additionally, species near the surface have a near-even adult sex ratio (ASR), whereas species from cave lakes have a female-biased ASR. We hypothesized (a) a decrease in sexual dimorphism from shallow subterranean habitats to cave lake species because of weaker sexual selection derived from changes in the ASR and (b) an increase in female body size in cave lakes because of stronger fecundity selection on account of oligotrophy, reduced predation, and weaker seasonality. We measured body size and two sexually dimorphic abdominal appendages for all 31 species and several behaviours related to male competition (activity, risk-taking, exploration) for 12 species. Species with an equal ASR that live close to the surface exhibited sexual dimorphism in all three morphological traits, but not in behaviour. The body size of females increased from the surface to cave lakes, but no such trend was observed in males. In cave lake species, males and females differed neither morphologically nor behaviourally. Our results are consistent with the possibility that sexual and fecundity selection covary across the three habitats, which indirectly and directly, respectively, shape the degree of sexual dimorphism in Niphargus species. [ABSTRACT FROM AUTHOR]

We evaluated whether the availability of male mates affected mating strategies in a predominantly socially monogamous passerine, the grass wren Cistothorus platensis. We used the natural variation in adult sex ratio (ASR) and a male removal manipulation to assess if social polygyny was more frequent when male mates were less abundant. We evaluated the potential costs that females paid when breeding with a polygynous male by assessing how males distributed parental care between nests and analysing four correlates of breeding success (clutch size, nestling body condition, number of fledglings, and nest fate). Social monogamy was predominant when males were abundant while social polygyny was more frequent when the ASR was female-biased. The removal of males was associated with a high rate of social polygyny. Social polygyny occurred when males annexed the territory of a neighbouring female or when an unpaired female settled within the territory of a mated male. We identified several potential costs of social polygyny to females. Secondary females (females that laid eggs after the primary female) produced nestlings of lower body condition during the first half of the breeding season, received less help in feeding the brood, and increased their parental contribution. We suggest that while intrasexual competition may constrain social polygyny when males are abundant, reduced female life expectancy combined with moderate costs of male desertion may facilitate social polygyny when potential male mates are not available. Significance statement: Social mating systems are thought to reflect the outcomes of optimal decisions made by females and males confronted with variation in the quality of territories and/or mates. However, demographic factors and life-history traits may constrain individual decisions. We studied how the availability of male mates might predispose grass wrens to social monogamy and constrain social polygyny. We found that social monogamy was predominant when males were more frequent while social polygyny increased when males were less abundant (female-biased adult sex ratio). Moreover, the removal of males in a plot early in the breeding season biased the adult sex ratio towards females and produced a polygyny rate comparable to rates in years when males were less abundant in other plots. Our results suggest that the abundance of male mates can affect individual mating decisions, by reducing male-male competition and/or the mating option for females. [ABSTRACT FROM AUTHOR]

Objectives: This paper starts from the unusual observation of the overrepresentation of females among adults in the cemetery of Bronze Age Shahr‐i Sokhta (Seistan, Iran) and explores the post marital residence pattern. By integrating taphonomical (skeletal preservation), anthropological (sex ratio [SR], sexual dimorphism, stress indicators, age at death), archeological (long distance trade indicators, habitation floor area, social role of women), and ancient DNA (heterozygosity levels in X chromosomes) data we test the hypothesis of post marital matrilocality in the site. Methods: We computed the SR (pelvis‐based sex determination) in a random unpublished adult sample from the cemetery of Shahr‐i Sokhta and in two samples previously published by other authors. We used comparative data on SR from: a large Supra Regional multi‐chronological sample of sites, n = 47, with 8808 adult sexed individuals, from Southern Europe, Egypt, Middle East, Southern Russia; a Regional Bronze Age sample of sites (n = 10) from Bactria Margiana and Indus Valley with 1324 adult sexed individuals. We estimated the heterozygosity levels in X chromosomes compared with the rest of the autosomes on the assumption that in a matrilocal society females should show lower variability than men. Results: Adult SR in a sample (n = 549) from Shahr‐i Sokhta is 70.5, the overrepresentation of females is shared with Regional Bronze Age sites from Bactria Margiana (SR = 72.09) and Indus Valley (SR = 67.54). On the contrary, in a larger Supra Regional multi‐chronological sample of sites, mean SR ranges between 112.7 (Bronze Age) and 163.1 (Middle Ages). Taphonomical and anthropological indicators do not explain the overrepresentation of female skeletons. Archeological indicators suggest a high social status of women and that the society was devoted to long range trade activities. heterozygosity levels in X chromosomes are in agreement with a matrilocal society. Conclusions: Indicators suggest that Bronze Age Shahr‐ì Sokhta was a matrilocal society and that long distance trade was an important economic factor producing an overrepresentation of adult female skeletons in the cemetery. [ABSTRACT FROM AUTHOR]

Males and females often exhibit different behaviours during mate acquisition, pair‐bonding and parenting, and a convenient label to characterize these behaviours is sex role. The diverse roles that male and female shorebirds (plovers, sandpipers and allies) exhibit in mating and parenting have played a key role in advancing mainstream theories in avian ecology and behavioural biology including sexual selection, sexual conflict and parental cooperation. Recent advances in shorebird research have also highlighted the significance of the social environment in driving sex role behaviours by linking the adult sex ratio with breeding behaviour and population demography. Here we review the key advances in sex role research using shorebirds as an ecological model system. We identify knowledge gaps and argue that shorebirds have untapped potential to accelerate diverse research fields including evolutionary genomics, movement ecology, social networks and environmental changes. Future studies of sex roles will benefit from individual‐based monitoring using advanced tracking technologies, and from multi‐team collaborations that are facilitated by standardized data collection methodologies across different species in the field. These advances will not only contribute to our understanding of reproductive strategies, but they will also have knock‐on effects on predicting population resilience to environmental changes and on prioritizing species for conservation. [ABSTRACT FROM AUTHOR]

In cooperatively breeding species, sexually mature individuals may delay natal dispersal and become subordinates, helping a dominant pair raise offspring. To understand how cooperative breeding evolved, it is important to determine the mechanisms leading to delayed dispersal. Adult sex ratio (ASR) variation may affect dispersal by limiting breeding vacancies available to the more abundant sex, and cooperative breeders often have a more biased ASR than noncooperative breeders. However, no studies of cooperative breeders have related ASR at both the local and population level with dispersal. Using the long-term Seychelles warbler, Acrocephalus sechellensis , data set, we tested the influence of population-wide and local ASR, and density, on natal dispersal of yearlings. Our ASR - density hypothesis predicts that the probability of natal dispersal is lower when the ASR is biased towards the sex of the focal individual, but only when the population density is high. Dispersal was associated with population density and population-wide ASR in males, but not in females; males were more likely to delay dispersal when ASR was male biased and density was high. Our findings illustrate a complex association between demographic factors and cooperative breeding and suggest that individuals alter their dispersal behaviour in response to the demographic composition of the population. • Natal dispersal is associated with ASR in a facultative cooperative breeder. • Male but not female yearlings disperse in response to population ASR and density. • Population-wide rather than local demographic composition predicted male dispersal. • Our study links demographic factors with behaviour that drives cooperative breeding. [ABSTRACT FROM AUTHOR]

We investigated ectoparasite diversity, interspecific infestation rates and host preference in roosting fruit bats, Eidolon helvum, from Bowen University, Southwest Nigeria. Fur of captured E. helvum were sampled monthly for ectoparasites from January 2021 to June 2022. We examined a total of 231 E. helvum and observed a significant female to male adult sex ratio (0.22:1); with 53.9% ectoparasitic infestation rate. We identified and enumerated the ectoparasite; and subjected its Cytochrome c oxidase subunit I (COI) gene to phylogenetic analysis with other nycteribiids. COI gene sequences obtained formed a distinct clade with other C. greeffi sequences. We recovered a total of 319 (149 female and 170 male) ectoparasites and observed a balanced C. greeffi female to male adult sex ratio of 0.88:1. Ectoparasitic sex distribution had no association with host sex and season. Prevalence was significantly higher during the wet season, but not between sexes of E. helvum. The intensity of infestation, 3.7 ± 0.4 individuals per fruit bat, was significantly higher during the wet season with a bimodal seasonal distribution. The strongly male-biased host adult sex ratio had no significant influence on C. greeffi metapopulation adult sex ratio.

Despite all efforts, long-term changes in the adult sex ratios of breeding duck populations are still unclear; this uncertainty is especially true for male-bias populations, which are often under the scrutiny of researchers lacking convenient results for the active protection of endangered species. Species with male-bias populations are usually strongly affected by a decline in population size that leads to a higher extinction risk. In this study, we examined our long-term data of the abundance of breeding populations in six duck species (Mallard Anas platyrhynchos, Gadwall Mareca strepera, Red-crested Pochard Netta rufina, Common Pochard Aythya ferina, Tufted Duck Aythya fuligula, and Common Goldeneye Bucephala clangula) from fishponds in South Bohemia, Czechia, between 2004 and 2022. This evidence was used to assess long-term changes in the adult sex ratio in these breeding populations and investigate the possible effects of the NAO index (North Atlantic Oscillation index) on them, indicating climate conditions in winter. We determined a long-term decrease of the proportion of females in the breeding season in two of the six examined species: Common Pochard and Red-crested Pochard, which is driven by the long-term increase in the number of males in contrast to the decreasing or stable number of females likely caused by different migration behaviours between females and males. In the case of Common Pochard, in breeding populations, we estimated 60–65% of males in the early 2000s rising to 75–80% in the early 2020s. However, we establish no significant effects linked to climate conditions of the previous winter in these species as a crucial cause of the changes of the proportion of females in the breeding population.

Traditional methods to study precopulatory mate guarding in Hyalella azteca (Saussure, 1858) have some limitations because precopulatory pairs are usually observed in isolation. This condition precludes researchers from testing the role of the social environment in the duration of precopulatory mate guarding. We present the use of a sampling method to investigate the influence of sex ratios on guarding success (the proportion of precopulatory pairs per trial) in mixed-sex groups of H. azteca. The presence/absence of precopulatory pairs over eight consecutive days, morning and afternoon samplings, was analysed as a series of successes (runs) to estimate the mean precopula duration. The non-random nature of the observed duration of the precopula was tested against a probabilistic distribution based on the set of all ways every possible duration may occur. While skewed sex ratios increased guarding success of the rarest sex, even sex ratios decreased it. The mean duration of observed precopulas (2.32 and 2.08 d for the morning and afternoon periods, respectively) is consistent with theoretical and empirical results, which confirms the accuracy and usefulness of the sampling method we used. The use of this method and the associated probabilistic analysis of the relevant data can enhance the study of the effect of biotic and abiotic factors on precopulatory mate guarding. It can also be used to test predictions derived from hypothesis concerned with mate guarding among crustaceans. [ABSTRACT FROM AUTHOR]

We investigated ectoparasite diversity, interspecific infestation rates and host preference in roosting fruit bats, Eidolon helvum , from Bowen University, Southwest Nigeria. Fur of captured E. helvum were sampled monthly for ectoparasites from January 2021 to June 2022. We examined a total of 231 E. helvum and observed a significant female to male adult sex ratio (0.22:1); with 53.9% ectoparasitic infestation rate. We identified and enumerated the ectoparasite; and subjected its Cytochrome c oxidase subunit I (COI) gene to phylogenetic analysis with other nycteribiids. COI gene sequences obtained formed a distinct clade with other C. greeffi sequences. We recovered a total of 319 (149 female and 170 male) ectoparasites and observed a balanced C. greeffi female to male adult sex ratio of 0.88:1. Ectoparasitic sex distribution had no association with host sex and season. Prevalence was significantly higher during the wet season, but not between sexes of E. helvum. The intensity of infestation, 3.7 ± 0.4 individuals per fruit bat, was significantly higher during the wet season with a bimodal seasonal distribution. The strongly male-biased host adult sex ratio had no significant influence on C. greeffi metapopulation adult sex ratio. [Display omitted] • The bat fly species, Cyclopodia greeffi, is ectoparasitic on Eidolon helvum. • Male biased adult sex ratio was observed for Eidolon helvum. • Prevalence and intensity of C. greeffi infestation showed seasonal significance. • There is no association between host and ectoparasite sex. [ABSTRACT FROM AUTHOR]

Managing for the effects of climate change on species whose populations are currently imperiled requires detailed knowledge of the relationship between their demographic rates and climate variables. We sought this information for the West Coast breeding population of snowy plover (Charadrius nivosus), which was federally listed as threatened in 1993 due to substantial declines in the numbers of plovers breeding along the coast and in the number of sites occupied for breeding. Snowy plovers employ a serially polygamous breeding system in which the male typically tends chicks to independence. This unusual breeding system is favored by male‐biased sex ratios in local populations. As part of a multispecies study of the effect of climate change on population growth, we used mark–capture models to examine climate drivers of adult survival for 1219 snowy plovers banded at Monterey Bay over 38 years and known to overwinter on the surrounding north‐central California outer coast. Nonclimate variables, including sex and unmeasured annual mortality risks (e.g., predator abundance), were the primary factors affecting adult survival. However, there is evidence that cold weather, particularly extended cold snaps with daily low temperatures below 2°C and daily high temperatures below 10°C, decreases overwinter survival. Exceptionally cold winters had a particularly strong effect on adult female plovers, contributing to the male‐biased adult sex ratios. Future winter climate on the north‐central California coast is projected to be generally warmer with fewer and shorter cold snaps. Reduced mortality from cold winter weather may mitigate other threats faced by plovers, such as anthropogenically enhanced predator populations, habitat loss, and accelerated sea level rise, while altering the adult sex ratio and potentially shifting the evolutionary landscape maintaining the plover's unusual breeding system. [ABSTRACT FROM AUTHOR]

Abstract Managing for the effects of climate change on species whose populations are currently imperiled requires detailed knowledge of the relationship between their demographic rates and climate variables. We sought this information for the West Coast breeding population of snowy plover (Charadrius nivosus), which was federally listed as threatened in 1993 due to substantial declines in the numbers of plovers breeding along the coast and in the number of sites occupied for breeding. Snowy plovers employ a serially polygamous breeding system in which the male typically tends chicks to independence. This unusual breeding system is favored by male‐biased sex ratios in local populations. As part of a multispecies study of the effect of climate change on population growth, we used mark–capture models to examine climate drivers of adult survival for 1219 snowy plovers banded at Monterey Bay over 38 years and known to overwinter on the surrounding north‐central California outer coast. Nonclimate variables, including sex and unmeasured annual mortality risks (e.g., predator abundance), were the primary factors affecting adult survival. However, there is evidence that cold weather, particularly extended cold snaps with daily low temperatures below 2°C and daily high temperatures below 10°C, decreases overwinter survival. Exceptionally cold winters had a particularly strong effect on adult female plovers, contributing to the male‐biased adult sex ratios. Future winter climate on the north‐central California coast is projected to be generally warmer with fewer and shorter cold snaps. Reduced mortality from cold winter weather may mitigate other threats faced by plovers, such as anthropogenically enhanced predator populations, habitat loss, and accelerated sea level rise, while altering the adult sex ratio and potentially shifting the evolutionary landscape maintaining the plover's unusual breeding system.

In recent years, researc in subjective well-being has found several ecological factors that may underpin societal differences in happiness. The adult sex ratio, the number of males relative to females in an environment, influences many behaviours in both humans and non-human animals. However, the possible influence of the sex ratio on subjective well-being has received little attention. I investigated the relationship between the adult sex ratio and subjective well-being in over 29,000 respondents from 133 regions of Europe. I find that women report lower subjective well-being in areas with more female-biased sex ratios, but males' well-being was unaffected. I did not find that the sex ratio influences the sex specific probability of marriage, or marriage rates overall. I also find that increased population density is associated with lower well-being. Drawing from sociological and evolutionary theories, I suggest that results may be due to females' decreased bargaining power in the dating market. [ABSTRACT FROM AUTHOR]

The origin and maintenance of sex differences in reproductive behavior (often labeled sex roles) have remained controversial topics, and recent meta-analyses and theoretical models have helped to elucidate the processes that generate diverse sex roles. We are glad to see that our study (Mokos et al., 2021) generated a healthy debate, and in agreement with recent commentaries (Janicke, 2024; Lehtonen & Parker, 2024) we call for a more comprehensive approach to understanding sex role evolution., (© The Author(s) 2024. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE) and European Society for Evolutionary Biology (ESEN).)

Adult sex ratio (ASR, the proportion of males in the adult population) is a central concept in population and evolutionary biology, and is also emerging as a major factor influencing mate choice, pair bonding and parental cooperation in both human and non-human societies. However, estimating ASR is fraught with difficulties stemming from the effects of spatial and temporal variation in the numbers of males and females, and detection/capture probabilities that differ between the sexes. Here, we critically evaluate methods for estimating ASR in wild animal populations, reviewing how recent statistical advances can be applied to handle some of these challenges. We review methods that directly account for detection differences between the sexes using counts of unmarked individuals (observed, trapped or killed) and counts of marked individuals using mark-recapture models. We review a third class of methods that do not directly sample the number of males and females, but instead estimate the sex ratio indirectly using relationships that emerge from demographic measures, such as survival, age structure, reproduction and assumed dynamics. We recommend that detection-based methods be used for estimating ASR in most situations, and point out that studies are needed that compare different ASR estimation methods and control for sex differences in dispersal.This article is part of the themed issue 'Adult sex ratios and reproductive decisions: a critical re-examination of sex differences in human and animal societies'.

Sexual violence against women is a global public health concern; yet, determining its patterning is still largely understudied. An excess of males has emerged as a central concern given that men are the primary perpetrators of violent behavior, particularly against women. However, it is increasingly unclear as to whether an excess or, rather, a shortage of men drives purported negative social outcomes. To address these conflicting expectations, we target data from the U.S. Census and the National Incident-Based Reporting System to explore the patterning of sexual violence against women across cities and counties in the United States. Through the use of generalized linear mixed models, we assess the role of adult sex ratio imbalance, along with measures of gender inequality, on sexual violence. Our results indicate mixed support for competing predictions. Violence does not simply increase by way of male surplus or shortage, but instead with increasing skew in the sex ratio. That is, balanced sex ratios exhibit the lowest rates of violence. However, rates of sexual violence against women increase more quickly with increasing male scarcity and are lowest at low levels of male excess (51%). Thus, our findings are particularly challenging to interpret from a 'more males = more violence' framework because violence increases more quickly with female excess and is rarest with a slight male bias in the population. We argue that these results highlight a need to target the specific types of violent behaviors expected to be motivated by partner availability, rather than overly simplistic predictions of male surfeit or deficit leading to an excess of violence. [ABSTRACT FROM AUTHOR]

Male-female agonism varies throughout the primate order with males often dominating females, especially in sexually dimorphic species. While intersexual agonism has been attributed to sexually coercive contexts, it can also occur for other reasons and intersexual dominance may be influenced by the adult sex ratio. If the proportion of males is high, certain males will regularly lose against other males. Loser-effects may then pave the way for some females to dominate these males, an effect that has been described in a few primate species. Here we investigated the frequency, general style, and context of agonism among gray langurs (Semnopithecus spp.). Data were collected at two study sites, at Jodhpur, India (one group), and at Ramnagar, Nepal (two groups). The adult sex ratio varied between 0.077 and 1.000 males to females. At both sites, data on agonistic interactions (aggression and submission) were collected in continuous focal animal and ad libitum sampling techniques during 1,945 contact hours (including 1,220 focal animal hours, total). Although aggression intensity was low, high directional consistency and the rare occurrence of counteraggression suggested a despotic dominance style, a likely prerequisite for dominance effects based on adult sex ratio. Aggression by females against males was very rare and mainly occurred in the defense of offspring. We found little evidence for partial female dominance regardless of adult sex ratio. In a few cases in which a female had a higher dominance index than a male, she did not dominate this male in dyadic encounters. Agonism by males directed at females occurred mainly in a feeding context while male policing and a sexual context were both rare. The latter was mostly restricted to females after they had harassed a sexual interaction or after they had behaved proceptively toward a male. Our study suggests that across species the effect of adult sex ratio on female dominance might be more variable than previously suggested. The fact that most agonism between males and females occurred over food identifies intersexual feeding competition as a new research avenue with potentially important consequences for existing ideas on the costs and benefits of group life and composition.

Sex roles describe sex differences in courtship, mate competition, social pair‐bonds and parental care. A key challenge is to identify associations among the components and the drivers of sex roles. Here, we investigate sex roles using data from over 1800 bird species. We found extensive variation and lability in proxies of sex roles, indicating remarkably independent evolution among sex role components. Climate and life history showed weak associations with sex roles. However, adult sex ratio is associated with sexual dimorphism, mating system and parental care, suggesting that social environment is central to explaining variation in sex roles among birds. Our results suggest that sex differences in reproductive behaviour are the result of diverse and idiosyncratic responses to selection. Further understanding of sex roles requires studies at the population level to test how local responses to ecology, life histories and mating opportunities drive processes that shape sex role variation among higher taxa. [ABSTRACT FROM AUTHOR]

Intersexual differences in life span (age at death) and aging (increase in mortality risk associated with functional deterioration) are widespread among animals, from nematodes to humans. Males often live shorter than females, but there is substantial unexplained variation among species and populations. Despite extensive research, it is poorly understood how life span differences between the sexes are modulated by an interplay among genetic, environmental and social factors.The goal of our study was to test how sex differences in life span and ageing are modulated by social and environmental factors, and by intrinsic differences between males and females.To disentangle the complex basis of sex differences in life span and aging, we combined comparative data from sex ratios in 367 natural populations of four species of African annual killifish with experimental results on sex differences in life span and aging from eight laboratory populations tested in treatments that varied social and environmental conditions.In the wild, females consistently outlived males. In captivity, sex‐specific mortality depended on social conditions. In social‐housed experimental groups, male‐biased mortality persisted in two aggressive species, but ceased in two placid species. When social and physical contacts were prevented by housing all fish individually, male‐biased mortality ceased in all four species. This outcome held across benign and challenging environmental conditions. Fitting demographic survival models revealed that increased baseline mortality was primarily responsible for a shorter male life span in social‐housing conditions. The timing and rate of aging were not different between the sexes. No marker of functional aging we recorded in our study (lipofuscin accumulation, proliferative changes in kidney and liver) differed between males and females, despite their previously confirmed association with functional aging in Nothobranchius killifish.We show that sex differences in life span and aging in killifish are driven by a combination of social and environmental conditions, rather than differential functional aging. They are primarily linked to sexual selection but precipitated through multiple processes (predation, social interference). This demonstrates how sex‐specific mortality varies among species even within an ecologically and evolutionary discrete lineage and explains how external factors mediate this difference. [ABSTRACT FROM AUTHOR]

Abstract Maturation (the age when organisms are physiologically capable of breeding) is one of the major life history traits that have pervasive implications for reproductive strategies, fitness, and population growth. Sex differences in maturation are common in nature, although the causes of such differences are not understood. Fisher and Lack proposed that delayed maturation in males is expected when males are under intense sexual selection, but their proposition has never been tested across a wide range of taxa. By using phylogenetic comparative analyses and the most comprehensive dataset to date, including 201 species from 59 avian families, we show that intense sexual selection on males (as indicated by polygamous mating and male‐skewed sexual size dimorphism) correlates with delayed maturation. We also show that the adult sex ratio (ASR), an indicator of the social environment, is associated with sex‐specific maturation because in species with a female‐skewed ASR, males experience later maturation. Phylogenetic path analyses suggest that adult sex ratio drives interspecific changes in the intensity of sexual selection which, in turn, influences maturation. These results are robust to alternative phylogenetic hypotheses and to potential life‐history confounds, and they provide the first comprehensive support of Fisher's and Lack's propositions. Importantly, our work suggests that both social environment and mate competition influence the evolution of a major life history trait, maturation.

The evolutionary history of sexual selection in the geologic past is poorly documented based on quantification, largely because of difficulty in sexing fossil specimens. Even such essential ecological parameters as adult sex ratio (ASR) and sexual size dimorphism (SSD) are rarely quantified, despite their implications for sexual selection. To enable their estimation, we propose a method for unbiased sex identification based on sexual shape dimorphism, using size-independent principal components of phenotypic data. We applied the method to test sexual selection in Keichousaurus hui, a Middle Triassic (about 237 Ma) sauropterygian with an unusually large sample size for a fossil reptile. Keichousaurus hui exhibited SSD biased towards males, as in the majority of extant reptiles, to a minor degree (sexual dimorphism index -0.087). The ASR is about 60% females, suggesting higher mortality of males over females. Both values support sexual selection of males in this species. The method may be applied to other fossil species. We also used the Gompertz allometric equation to study the sexual shape dimorphism of K. hui and found that two sexes had largely homogeneous phenotypes at birth except in the humeral width, contrary to previous suggestions derived from the standard allometric equation.

Abstract A sex ratio is one of the most basic demographic estimates produced because it is easy to collect and provides deeper insight into population dynamics for the species under consideration. For inconsistently or intermittently breeding species, the breeding sex ratio (BSR) and adult sex ratio (ASR), both reported as the proportion of males, can be quite different. The entire adult population of some wide‐ranging species may never be present and capable of being sampled in the same time and place. We explore equations to indirectly estimate ASRs and annual abundance estimates from annual surveys of BSRs. We sampled Atlantic sturgeon (Acipenser oxyrinchus oxyrinchus) from 2013 through 2019 and implanted acoustic transmitters during those sampling periods. The BSRs calculated during capture from 2015 through 2019 were 0.65, 0.75, 0.69, 0.75, and 0.64 each year. Relying on telemetry detections from the lowest potential spawning region, the expected BSRs in the same years were 0.64, 0.74, 0.67, 0.69, and 0.60, suggesting telemetry is a reliable and passive way to estimate BSR. The BSRs were used to indirectly estimate ASR to be approximately 0.51 (95% confidence limits of 0.43–0.58). Estimates of annual abundance derived through sex ratios matched previously published mark–recapture estimates of the same breeding population, but provide additional detail on abundances of each sex. For populations where BSR is more accessible, ASR and abundance estimates can be estimated with capture data and acoustic telemetry.

Males and females follow distinct life‐history strategies that have co‐evolved with several sex‐specific traits. Higher investment into parental investment (PI) demands an increased lifespan. Thus, resource allocation toward an efficient immune system is mandatory. In contrast, resources allocated toward secondary sexual signals (ornamentation) may negatively correlate with investment into immunity and ultimately result in a shorter lifespan. Previous studies have addressed how resource allocation toward single sex‐specific traits impacts lifetime reproductive success (LRS). However, the trade‐offs between diverse sex‐specific characteristics and their impact on LRS remain largely unassessed impeding our understanding of life‐history evolution. We have designed a theoretical framework (informed by experimental data and evolutionary genetics) that explores the effects of multiple sex‐specific traits and assessed how they influence LRS. From the individual sex‐specific traits, we inferred the consequences at the population level by evaluating adult sex ratios (ASR). Our theory implies that sex‐specific resource allocation toward the assessed traits resulted in a biased ASR. Our model focuses on the impact of PI, ornamentation, and immunity as causal to biased ASR. The framework developed herein can be employed to understand the combined impact of diverse sex‐specific traits on the LRS and the eventual population dynamics of particular model systems. [ABSTRACT FROM AUTHOR]

A sex ratio is one of the most basic demographic estimates produced because it is easy to collect and provides deeper insight into population dynamics for the species under consideration. For inconsistently or intermittently breeding species, the breeding sex ratio (BSR) and adult sex ratio (ASR), both reported as the proportion of males, can be quite different. The entire adult population of some wide‐ranging species may never be present and capable of being sampled in the same time and place. We explore equations to indirectly estimate ASRs and annual abundance estimates from annual surveys of BSRs. We sampled Atlantic sturgeon (Acipenser oxyrinchus oxyrinchus) from 2013 through 2019 and implanted acoustic transmitters during those sampling periods. The BSRs calculated during capture from 2015 through 2019 were 0.65, 0.75, 0.69, 0.75, and 0.64 each year. Relying on telemetry detections from the lowest potential spawning region, the expected BSRs in the same years were 0.64, 0.74, 0.67, 0.69, and 0.60, suggesting telemetry is a reliable and passive way to estimate BSR. The BSRs were used to indirectly estimate ASR to be approximately 0.51 (95% confidence limits of 0.43–0.58). Estimates of annual abundance derived through sex ratios matched previously published mark–recapture estimates of the same breeding population, but provide additional detail on abundances of each sex. For populations where BSR is more accessible, ASR and abundance estimates can be estimated with capture data and acoustic telemetry. [ABSTRACT FROM AUTHOR]

Sexual violence against women is a global public health concern; yet, determining its patterning is still largely understudied. An excess of males has emerged as a central concern given that men are the primary perpetrators of violent behavior, particularly against women. However, it is increasingly unclear as to whether an excess or, rather, a shortage of men drives purported negative social outcomes. To address these conflicting expectations, we target data from the U.S. Census and the National Incident-Based Reporting System to explore the patterning of sexual violence against women across cities and counties in the United States. Through the use of generalized linear mixed models, we assess the role of adult sex ratio imbalance, along with measures of gender inequality, on sexual violence. Our results indicate mixed support for competing predictions. Violence does not simply increase by way of male surplus or shortage, but instead with increasing skew in the sex ratio. That is, balanced sex ratios exhibit the lowest rates of violence. However, rates of sexual violence against women increase more quickly with increasing male scarcity and are lowest at low levels of male excess (51%). Thus, our findings are particularly challenging to interpret from a ‘more males = more violence’ framework because violence increases more quickly with female excess and is rarest with a slight male bias in the population. We argue that these results highlight a need to target the specific types of violent behaviors expected to be motivated by partner availability, rather than overly simplistic predictions of male surfeit or deficit leading to an excess of violence.

The relative number of adult males to females in a population, the adult sex ratio (ASR), is an important demographic variable in populations, which plays a role in, for instance, mating systems and viability of populations. The origins of variation in ASR between populations are often unclear. A skewed ASR in a local population may result from a skewed sex ratio among fledglings, sex differences in survival or dispersal or a combination of these. Meta-analyses show that ASRs across bird species are mostly male-biased and that male-biased survival is the main contributor to this bias. Here, we examine the origins of a female-biased ASR in a small population (3–24 breeding females in 2007–2017) of a passerine, the Northern Wheatear Oenanthe oenanthe. Because 95% of all fledglings were sexed, 88% of the individuals were colour-banded and because this species is highly philopatric, we could accurately examine: (1) fledgling sex ratio, (2) sex-specific first year and adult apparent survival and (3) sex-specific dispersal. It appeared that survival was not sex-biased for either age cohort but that fledgling sex ratio was male-biased, especially so among late broods. In addition, natal dispersal out of the breeding population was likely male-biased, in contrast to the general pattern of female-bias in avian dispersal. So, perhaps females are more common in our relict population solely because more males disperse from the population than females, without being compensated by male-biased fledgling sex ratio or survival and without compensation from immigrating males. Likely, the population sex ratio (including both adults and young) at the end of the breeding season is male-biased, because of a pronounced male-biased sex ratio among fledglings from late nests. Perhaps this male-bias induces prospecting young males to settle elsewhere to avoid competition next year, but at the same time convinces prospecting females to settle particularly in this area. [ABSTRACT FROM AUTHOR]

Variation in female mating performance can affect the direction and rate of evolution through sexual selection. The social environment determines the availability of mates and the competitive situation, and hence can influence mating strategies. However, these effects have to be considered within time constraints. Here we show that female sand gobies exposed to same-sex competitors (female-biased adult sex ratio, ASR) for a week before having physical access to males (i.e., a delayed male access; referred as prior exposure treatment) were more actively associated with the preferred male and took faster spawning decisions. However, these females mated more frequently with males with traits that did not ensure high egg survival. On the other extreme, females exposed to low same sex competition (male-biased ASR) simultaneously to the mate choice (i.e., an immediate access to males; referred as simultaneous exposure treatment) took more time to make their spawning decision. They also associated and spawned more often with males with good parental skills (i.e., males exhibiting more egg fanning behaviour). These male traits are associated with higher survival and better development of eggs. Our results provide experimental support for a trade-off between mate choice accuracy and speed, which depended on mating competition. [ABSTRACT FROM AUTHOR]

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Group living may entail local resource competition (LRC) which can be reduced if the birth sex ratio (BSR) is biased towards members of the dispersing sex who leave the group and no longer compete locally with kin. In primates, the predicted relationship between dispersal and BSR is generally supported although data for female dispersal species are rare and primarily available from captivity. Here, we present BSR data for Phayre's leaf monkeys ( Trachypithecus phayrei crepusculus ) at the Phu Khieo Wildlife Sanctuary, Thailand ( N = 104). In this population, nearly all natal females dispersed, while natal males stayed or formed new groups nearby. The slower reproductive rate in larger groups suggests that food can be a limiting resource. In accordance with LRC, significantly more females than males were born (BSR 0.404 males/all births) thus reducing future competition with kin. This bias was similar in 2-year-olds (no sex-differential mortality). It became stronger in adults, supporting our impression of particularly fierce competition among males. To better evaluate the importance of BSR, more studies should report sex ratios throughout the life span, and more data for female dispersal primates need to be collected, ideally for multiple groups of different sizes and for several years.

The adult sex ratio (ASR) is important within ecology due to its predicted effects on behaviour, demography and evolution, but research examining the causes and consequences of ASR bias have lagged behind the studies of sex ratios at earlier life stages. Although ungulate ASR is relatively well‐studied, exceptions to the usual female‐biased ASR challenge our understanding of the underlying drivers of biased ASR and provide an opportunity to better understand its consequences.Some feral ungulate populations, including multiple horse populations, exhibit unusually male‐biased ASR. For example, research suggests that the feral horse Equus ferus caballus population on Sable Island, Nova Scotia, Canada may exhibit a male‐biased ASR. Such exceptions to the rule provide a valuable opportunity to reveal the contributions of environmental context and trait differences to ASR bias.We aimed to test for bias in Sable Island horse ASR, identify the demographic drivers of bias, and explore its demographic and social consequences. To do this, we used life history, movement and group membership information for hundreds of horses followed through a long‐term individual‐based study between 2007 and 2018.Sable Island horse ASR is male biased and this skew has increased over time, reaching 62% male in 2018. Our life table response experiment suggested that ASR skew was driven predominantly by male‐biased adult survival. Further analyses pointed to sex‐biased survival being driven by reduced female survival post‐reproduction. Male‐biased ASR was associated with reduced harem sizes, an increase in the number of social groups on the island, and reduced reproduction in young females.Our results support the idea that male‐biased ASR in feral ungulate populations may be caused by a combination of high population density and high reproductive output. We suggest that female‐biased mortality may be caused by females continuing to reproduce at high density, and thus being more susceptible to resource shortages. Thus, our results highlight the strong context dependence of ASR. Furthermore, our work indicates the potential for ASR to substantially alter a population's social organisation. Such changes in social structure could have knock‐on consequences for demography by altering the formation/stability of social relationships, or competition for matings. [ABSTRACT FROM AUTHOR]

Maturation (the age when organisms are physiologically capable of breeding) is one of the major life history traits that have pervasive implications for reproductive strategies, fitness, and population growth. Sex differences in maturation are common in nature, although the causes of such differences are not understood. Fisher and Lack proposed that delayed maturation in males is expected when males are under intense sexual selection, but their proposition has never been tested across a wide range of taxa. By using phylogenetic comparative analyses and the most comprehensive dataset to date, including 201 species from 59 avian families, we show that intense sexual selection on males (as indicated by polygamous mating and male‐skewed sexual size dimorphism) correlates with delayed maturation. We also show that the adult sex ratio (ASR), an indicator of the social environment, is associated with sex‐specific maturation because in species with a female‐skewed ASR, males experience later maturation. Phylogenetic path analyses suggest that adult sex ratio drives interspecific changes in the intensity of sexual selection which, in turn, influences maturation. These results are robust to alternative phylogenetic hypotheses and to potential life‐history confounds, and they provide the first comprehensive support of Fisher's and Lack's propositions. Importantly, our work suggests that both social environment and mate competition influence the evolution of a major life history trait, maturation. [ABSTRACT FROM AUTHOR]

Recently, it was pointed out that classic models for the evolution of anisogamy do not take into account the possibility of parthenogenetic reproduction, even though sex is facultative in many relevant taxa (e.g., algae) that harbour both anisogamous and isogamous species. Here, we complement this recent analysis with an approach where we assume that the relationship between progeny size and its survival may differ between parthenogenetically and sexually produced progeny, favouring either the former or the latter. We show that previous findings that parthenogenesis can stabilise isogamy relative to the obligate sex case, extend to our scenarios. We additionally investigate two different ways for one mating type to take over the entire population. First, parthenogenesis can lead to biased sex ratios that are sufficiently extreme that one type can displace the other, leading to de facto asexuality for the remaining type that now lacks partners to fuse with. This process involves positive feedback: microgametes, being numerous, lack opportunities for syngamy, and should they proliferate parthenogenetically, the next generation makes this asexual route even more prominent for microgametes. Second, we consider mutations to strict asexuality in producers of micro- or macrogametes, and show that the prospects of asexual invasion depend strongly on the mating type in which the mutation arises. Perhaps most interestingly, we also find scenarios in which parthenogens have an intrinsic survival advantage yet facultatively sexual isogamous populations are robust to the invasion of asexuals, despite us assuming no genetic benefits of recombination. Here, equal contribution from both mating types to zygotes that are sufficiently well provisioned can outweigh the additional costs associated with syngamy.

Sex ratio variation has important consequences for population dynamics and viability. Adult sex ratio is male biased in many duck species, and data from wintering areas suggest male bias has been increasing in some European duck species. However, the reasons for changes in sex ratios remain poorly understood, and information on sex ratio variation among breeding ducks is limited. We studied long-term changes in breeding sex ratios and breeding numbers in Common Pochard Aythya ferina and Tufted Duck Aythya fuligula in southern Finland, the former species being currently assessed as critically endangered and the latter as endangered in Finland. In addition, we tested the hypothesis that between-year variation in breeding sex ratios is affected by the severity of weather conditions in the wintering areas. The proportion of females among breeding Tufted Ducks decreased from 42.9% in 1951–1970 to 36.9% in 1996–2015, while no statistically significant change was observed in Common Pochard (41.8% and 39.5%, respectively). Both species showed a decline in breeding numbers over the same period. Severity of the preceding winter did not affect the proportion of females in the breeding area. [ABSTRACT FROM AUTHOR]

A biased adult sex ratio (ASR) can influence cooperative breeding behavior if the bias limits mating opportunities for the more abundant sex. We tested predictions associated with the ASR-cooperation hypothesis in the brown-headed nuthatch (Sitta pusilla). We manipulated ASR by cross-fostering known-sex nestlings within 2 large (≥100 ha) experimental plots for 5 years using a crossover design where each plot received an opposing male- or female-biased treatment for 2 consecutive years. A year with no manipulations followed before the bias was reversed on each plot for 2 additional years. Variation in ASR (adult males/total adults) was pronounced compared to background proportions (0.55) and ranged from a female bias in female-biased plots (0.47) to a strong male bias in male-biased plots (0.71). Sex ratios during the postbreeding period ranged more broadly (0.33 in female-biased plots vs. 0.74 in male-biased plots). Territory densities did not change significantly and allowed 6 predictions to be assessed. Consistent with predictions, the prevalence of cooperative breeding groups doubled under male-biased treatments and large cooperative groups appeared (≥2 male helpers vs. the single male helper most common prior to the experiment). These changes occurred despite increased dispersal of cross-fostered males in male-biased plots. Most juvenile females dispersed, but, consistent with predictions, the prevalence of female helpers increased under female-biased treatments. Manipulations did not alter the sex of nestlings produced nor extend the time that males served as helpers. Taken collectively, results support the ASR-cooperation hypothesis and the role that mate limitations play in cooperative breeding behavior. [ABSTRACT FROM AUTHOR]

The adult sex ratio (ASR) is a crucial component of the ecological and evolutionary forces shaping the dynamics of a population. Although in many declining populations ASRs have been reported to be skewed, empirical studies exploring the demographic factors shaping ASRs are still rare. In this study of the socially monogamous and sexually dimorphic Black-tailed Godwit (Limosa limosa limosa), we aim to evaluate the sex ratio of chicks at hatch and the subsequent sex-specific survival differences [ABSTRACT FROM AUTHOR]

In Focus:Heinsohn, R., Olah, G., Webb, M., Peakall, R., & Stojanovic, D. (2019). Sex ratio bias and shared paternity reduce individual fitness and population viability in a critically endangered parrot. Journal of Animal Ecology, 88, 502–510. While the effects of variation in the sex ratio of offspring have been thoroughly explored over the last century, the sex ratio of adults has received far less attention. A paper by Heinsohn and colleagues in this issue shows that changes in the adult sex ratio can have striking effects on mating systems, reproductive success and population viability. These impacts are all the more dramatic because they occur in a critically endangered species, the swift parrot, Lathamus discolor, of Tasmania. This species suffers heavy predation from the introduced sugar glider, Petaurus breviceps, which kills nesting females and their clutches, resulting in strongly male‐biased adult sex ratios. The authors combined demographic and genetic data to show that, at sites with heavier predation, the remaining females were more likely to mate with multiple males. This shift in the mating system also led to lower overall nesting success, with fewer chicks fledged per nest at sites with higher levels of mixed paternity. Population viability models based on these data predicted steep population declines, with models using the highest observed rates of mixed paternity showing the sharpest declines. These results demonstrate that changes in the adult sex ratio can have far‐reaching impacts, including on the fitness of populations themselves. This In Focus highlights the striking effects of predator‐induced changes in mating system on the population fitness of an endangered parrot described in Heinsohn et al. (). [ABSTRACT FROM AUTHOR]

Sex‐biased mortality can lead to altered adult sex ratios (ASRs), which may in turn lead to harassment and lower fitness of the rarer sex and changes in the mating system. Female critically endangered swift parrots (Lathamus discolor) suffer high predation while nesting due to an introduced mammalian predator, the sugar glider (Petaurus breviceps). High predation on females is causing severe population decline alongside strongly biased adult sex ratios (≥73% male).Our 6‐year study showed that 50.5% of critically endangered swift parrot nests had shared paternity although the birds remained socially monogamous. Shared paternity increased significantly with the local rate of predation on breeding females, suggesting that rates of shared paternity increased when the ASR became more biased.Nests that were not predated produced fewer fledglings as the local ASR became more male‐biased possibly due to higher interference during nesting from unpaired males.Population viability analyses showed that part of the predicted decline in the swift parrot population is due to reduced reproductive success when paternity is shared. The models predicted that the population would decline by 89.4% over three generations if the birds maintained the lowest observed rate of shared paternity. This compares with predicted population reductions of 92.1–94.9% under higher rates of shared paternity.We conclude that biases in the ASR, in this case caused by sex‐specific predation from an introduced predator, can lead to changes in the mating system and negative impacts on both individual fitness and long‐term population viability. This paper offers rare insights into the effects of skewed adult sex ratios due to anthropogenic factors. Sex‐specific predation in swift parrots leads to strongly male‐biased adult sex ratios and unusually high rates of shared paternity (50.5%). Shared paternity is costly and has negative impacts on fitness and population viability. [ABSTRACT FROM AUTHOR]

Abstract: Sex‐biased dispersal is common in vertebrates, although the ecological and evolutionary causes of sex differences in dispersal are debated. Here, we investigate sex differences in both natal and breeding dispersal distances using a large dataset on birds including 86 species from 41 families. Using phylogenetic comparative analyses, we investigate whether sex‐biased natal and breeding dispersal are associated with sexual selection, parental sex roles, adult sex ratio (ASR), or adult mortality. We show that neither the intensity of sexual selection, nor the extent of sex bias in parental care was associated with sex‐biased natal or breeding dispersal. However, breeding dispersal was related to the social environment since male‐biased ASRs were associated with female‐biased breeding dispersal. Male‐biased ASRs were associated with female‐biased breeding dispersal. Sex bias in adult mortality was not consistently related to sex‐biased breeding dispersal. These results may indicate that the rare sex has a stronger tendency to disperse in order to find new mating opportunities. Alternatively, higher mortality of the more dispersive sex could account for biased ASRs, although our results do not give a strong support to this explanation. Whichever is the case, our findings improve our understanding of the causes and consequences of sex‐biased dispersal. Since the direction of causality is not yet known, we call for future studies to identify the causal relationships linking mortality, dispersal, and ASR. [ABSTRACT FROM AUTHOR]

Abstract: Adult sex ratio (ASR) plays a pivotal role in many ecological processes, such as population dynamics in time and space, extinction risk and reproductive success, but few patterns in the variability in ASR have been found across species. Due to the socio‐spatial structure and polygynous mating system found in felids, ASR is expected to be female‐biased in felids in a finite area; however, ASR is also commonly found to be either balanced or male‐biased in this taxonomic group. Therefore, investigating ecological correlates of the pattern of ASR variability in felids is of prime importance both for a better understanding of evolutionary ecology and for a more evidence‐based conservation of these threatened taxa. The aim of this review was to investigate whether the expected female‐biased ASR occurs in felids, and whether ASR is influenced by the size of the study area, the species’ body mass and the population density, taking into account the species‐specific home‐range requirements. We reviewed the camera‐trapping literature on felids to build a large global multi‐species data set on ASR variation. We used general linear mixed models to investigate ecological correlates of ASR. In small study areas, the ASR was balanced for all species, but in large areas the ASR was female‐biased for large and medium cats and male‐biased for small cats; sampling designs that are unsuitable for small cats (due to ‘gaps’ within the trapping grid) might explain this latter finding. Moreover, in large study areas, an increase in cat density was associated with more females, whereas in small areas the reverse was true. Our study provides a scientifically sound background for management and conservation actions of these flagship‐threatened taxa of carnivores, by indicating the importance of the study area on ASR in felids. Large areas can promote the recovery of large and medium cats, as more females can be hosted. [ABSTRACT FROM AUTHOR]

Our understanding of sexual selection has greatly improved during the last decades. The focus is no longer solely on males, but also on how female competition and male mate choice shape ornamentation and other sexually selected traits in females. At the same time, the focus has shifted from documenting sexual selection to exploring variation and spatiotemporal dynamics of sexual selection, and their evolutionary consequences. Here, I review insights from a model system with exceptionally dynamic sexual selection, the two-spotted goby fish Gobiusculus flavescens. The species displays a complete reversal of sex roles over a 3-month breeding season. The reversal is driven by a dramatic change in the operational sex ratio, which is heavily male-biased at the start of the season and heavily female-biased late in the season. Early in the season, breeding-ready males outnumber mature females, causing males to be highly competitive, and leading to sexual selection on males. Late in the season, mating-ready females are in excess, engage more in courtship and aggression than males, and rarely reject mating opportunities. With typically many females simultaneously courting available males late in the season, males become selective and prefer more colorful females. This variable sexual selection regime likely explains why both male and female G. flavescens have ornamental colors. The G. flavescens model system reveals that sexual behavior and sexual selection can be astonishingly dynamic in response to short-term fluctuations in mating competition. Future work should explore whether sexual selection is equally dynamic on a spatial scale, and related spatiotemporal dynamics. [ABSTRACT FROM AUTHOR]

Theoretical models suggest that adult sex ratio (ASR) and population density are expected to influence parental roles by reducing the mating opportunities of the commoner sex and by changing the intensity of sperm competition, although experimental evidence for these predictions is sparse. In biparental species with a high risk of extrapair paternity and consecutive egg laying over the breeding period, males are expected to reduce their parental investment and to spend more time on mate guarding if male density is high, to maximize their fitness. We conducted a field experiment to test this hypothesis in Lethrus apterus, a flightless biparental beetle species from the Geotrupidae family. Using seminatural enclosures, we assigned individuals to nine treatment groups differing in adult sex ratio (three levels) and individual density (three levels) using a full factorial experimental design. Nest attendance and parental provisioning (i.e. collecting and transporting leaves to the nest) were recorded as well as the number, size and sex ratio of the offspring. We found that as the level of male–male competition increased, generated either by the increased density of individuals or by the male-biased sex ratio, pairs showed higher nest attendance and collected fewer leaves. Male-biased groups also produced fewer offspring under high and low densities indicating a possible conflict of interest between the sexes over paternity and brood size. These results support the increased paternity assurance hypothesis under a high level of intrasexual competition. [ABSTRACT FROM AUTHOR]

Evidence from animal species indicates that a male-biased adult sex ratio (ASR) can lead to higher levels of male parental investment and that there is heterogeneity in behavioural responses to mate scarcity depending on mate value. In humans, however, there is little consistent evidence of the effect of the ASR on pair-bond stability and parental investment and even less of how it varies by an individual's mate value. In this paper we use detailed census data from Northern Ireland to test the association between the ASR and pair-bond stability and parental investment by social status (education and social class) as a proxy for mate value. We find evidence that female, but not male, cohabitation is associated with the ASR. In female-biased areas women with low education are less likely to be in a stable pair-bond than highly educated women, but in male-biased areas women with the lowest education are as likely to be in a stable pair-bond as their most highly educated peers. For both sexes risk of separation is greater at female-biased sex ratios. Lastly, our data show a weak relationship between parental investment and the ASR that depends on social class. We discuss these results in the light of recent reformulations of parental investment theory. [ABSTRACT FROM AUTHOR]

Sex reversals whereby individuals of one genetic sex develop the phenotype of the opposite sex occur in ectothermic vertebrateswith genetic sex-determination systems that are sensitive to extreme temperatures during sexual differentiation. Recent rises in global temperatures have led researchers to predict that sex reversals will become more common, resulting in the distortion of many populations' sex ratios. However, it is unclear whether susceptibility to climate-driven sex-ratio shifts depends on the type of sex determination that varies across species. First, we show here using individual-based theoretical models that XX/XY (male-heterogametic) and ZZ/ZW(female-heterogametic) sex-determination systems can respond differentially to temperature-induced sex reversals. Interestingly, the impacts of climate warming on adult sex ratio (ASR) depend on the effects of both genotypic and phenotypic sex on survival and reproduction. Second, we analyse the temporal changes of ASR in natural amphibian populations using data from the literature, and find that ASR shifted towards males in ZZ/ZW species over the past 60 years, but did not change significantly in XX/XY species. Our results highlight the fact that we need a better understanding of the interactions between genetic and environmental sex-determining mechanisms to predict the responses of ectotherms to climate change and the associated extinction risks. [ABSTRACT FROM AUTHOR]

Cooperative breeding is a form of breeding system where in addition to a core breeding pair, one or more usually non-breeding individuals provide offspring care. Cooperative breeding is widespread in birds, but its origin and maintenance in contemporary populations are debated. Although deviations in adult sex ratio (ASR, the proportion of males in the adult population) have been hypothesized to influence the occurrence of cooperative breeding because of the resulting surplus of one sex and limited availability of breeding partners, this hypothesis has not been tested across a wide range of taxa. By using data from 188 bird species and phylogenetically controlled analyses, we show that cooperatively breeding species have more male-biased ASRs than non-cooperative species. Importantly, ASR predicts helper sex ratio: in species with more male-biased ASR, helper sex ratio is also more male biased. We also show that offspring sex ratios do not predict ASRs, so that the skewed ASRs emerge during the period when individuals aim to obtain a breeding position or later during adulthood. In line with this result, we found that ASR (among both cooperatively and non-cooperatively breeding species) is inversely related to sex bias in dispersal distance, suggesting that the cost of dispersal is more severe for the further-dispersing sex. As females usually disperse further in birds, this explains the generally male-biased ASR, and in combination with benefits of philopatry for males, this probably explains why ASR is more biased in cooperatively breeding species. Taken together, our results suggest that a sex bias in helping in cooperatively breeding species relates to biased ASRs. We propose that this relationship is driven by sex-specific costs and benefits of dispersal and helping, as well as other demographic factors. Future phylogenetic comparative and experimental work is needed to establish how this relationship emerges. [ABSTRACT FROM AUTHOR]

Theoretical models and empirical studies in various taxa have identified important links between variation in sex roles and the number of adult males and females (adult sex ratio (ASR)) in a population. In this review, I examine these relationships in non-human primates. Because most existing theoretical models of the evolution of sex roles focus on the evolutionary origins of sex-biased behaviour, they offer only a general scaffold for predicting variation in sex roles among and within species. I argue that studies examining sex role variation at these more specific levels need to take social organization into account to identify meaningful levels for the measurement of ASR and to account for the fact that ASR and sex roles mutually influence each other. Moreover, taxon-specific life-history traits can constrain sex role flexibility and impact the operational sex ratio (OSR) by specifying the minimum length of female time outs from reproduction. Using examples from the primate literature, I highlight practical problems in estimating ASR and OSR. I then argue that interspecific variation in the occurrence of indirect forms of paternal care might indeed be linked to variation in ASR. Some studies also indicate that female aggression and bonding, as well as components of intersexual relationships, are sensitive to variation in ASR. Thus, links between primate sex roles and sex ratios merit further study, and such studies could prompt the development of more specific theoretical models that make realistic assumptions about taxon-specific life history and social organization. [ABSTRACT FROM AUTHOR]

Small populations are susceptible to high genetic loads and random fluctuations in birth and death rates. While these selective forces can adversely affect their viability, small populations persist across taxa. Here, we investigate the resilience of small groups to demographic uncertainty, and specifically to fluctuations in adult sex ratio (ASR), partner availability and dispersal patterns. Using 25 years of demographic data for two Savannah Pume' groups of South American hunter-gatherers, we show that in small human populations: (i) ASRs fluctuate substantially from year to year, but do not consistently trend in a sex-biased direction; (ii) the primary driver of local variation in partner availability is stochasticity in the sex ratio at maturity; and (iii) dispersal outside of the group is an important behavioural means to mediate locally constrained mating options. To then simulate conditions under which dispersal outside of the local group may have evolved, we develop two mathematical models. Model results predict that if the ASR is biased, the globally rarer sex should disperse. The model's utility is then evaluated by applying our empirical data to this central prediction. The results are consistent with the observed hunter-gatherer pattern of variation in the sex that disperses. Together, these findings offer an alternative explanation to resource provisioning for the evolution of traits central to human sociality (e.g. flexible dispersal, bilocal post-marital residence and cooperation across local groups). We argue that in small populations, looking outside of one's local group is necessary to find a mate and that, motivated by ASR imbalance, the alliances formed to facilitate the movement of partners are an important foundation for the human-typical pattern of network formation across local groups. [ABSTRACT FROM AUTHOR]

It is increasingly recognized that the relative proportion of potential mates to competitors in a population impacts a range of sex-specific behaviours and in particular mating and reproduction. However, while the adult sex ratio (ASR) has long been recognized as an important link between demography and behaviour, this relationship remains understudied. Here, we introduce the first inter-disciplinary collection of research on the causes and consequences of variation in the ASR in human and animal societies. This important topic is relevant to a wide audience of both social and biological scientists due to the central role that the relative number of males to females in a population plays for the evolution of, and contemporary variation in, sex roles across groups, species and higher taxa. The articles in this theme issue cover research on ASR across a variety of taxa and topics. They offer critical re-evaluations of theoretical foundations within both evolutionary and non-evolutionary fields, and propose innovative methodological approaches, present new empirical examples of behavioural consequences of ASR variation and reveal that the ASR plays a major role in determining population viability, especially in small populations and species with labile sex determination. This introductory paper puts the contributions of the theme issue into a broader context, identifies general trends across the literature and formulates directions for future research. [ABSTRACT FROM AUTHOR]