Critical Re-examination Of Sex Differences Research Articles

Adult sex ratios and reproductive strategies: a critical re-examination of sex differences in human and animal societies.

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.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'.

Adult sex ratios and partner scarcity among hunter-gatherers: implications for dispersal patterns and the evolution of human sociality.

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 Pumé 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.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'.

Evolution of male strategies with sex-ratio-dependent pay-offs: connecting pair bonds with grandmothering.

Men's provisioning of mates and offspring has been central to ideas about human evolution because paternal provisioning is absent in our closest evolutionary cousins, the great apes, and is widely assumed to result in pair bonding, which distinguishes us from them. Yet mathematical modelling has shown that paternal care does not readily spread in populations where competition for multiple mates is the common male strategy. Here we add to models that point to the mating sex ratio as an explanation for pairing as pay-offs to mate guarding rise with a male-biased sex ratio. This is of interest for human evolution because our grandmothering life history shifts the mating sex ratio from female- to male-biased. Using a difference equation model, we explore the relative pay-offs for three competing male strategies (dependant care, multiple mating, mate guarding) in response to changing adult sex ratios. When fertile females are abundant, multiple mating prevails. As they become scarce, mate guarding triumphs. The threshold for this shift depends on guarding efficiency. Combined with mating sex ratios of hunter-gatherer and chimpanzee populations, these results strengthen the hypothesis that the evolution of our grandmothering life history propelled the shift to pair bonding in the human lineage.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'.

The potential pitfalls of studying adult sex ratios at aggregate levels in humans.

Human adult sex ratios have been studied extensively across the biological and social sciences. While several studies have examined adult sex ratio effects in a multilevel perspective, many studies have focused on effects at an aggregated level only. In this paper, we review some key issues relating to such analyses. We address not only nation-level analyses, but also aggregation at lower levels, to investigate whether these issues extend to lower levels of aggregation. We illustrate these issues with novel databases covering a broad range of variables. Specifically, we discuss distributional issues with aggregated measures of adult sex ratio, significance testing, and statistical non-independence when using aggregate data. Firstly, we show that there are severe distributional issues with national adult sex ratio, such as extreme cases. Secondly, we demonstrate that many 'meaningless' variables are significantly correlated with adult sex ratio (e.g. the max. elevation level correlates with sex ratio at US state level). Finally, we re-examine associations between adult sex ratios and teenage fertility and find no robust evidence for an association at the aggregate level. Our review highlights the potential issues of using aggregate data on adult sex ratios to test hypotheses from an evolutionary perspective in humans.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'.

Sex roles and adult sex ratios: insights from mammalian biology and consequences for primate behaviour.

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 inter-sexual 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.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'.

Social determinants of adult sex ratios and racial/ethnic disparities in transmission of HIV and other sexually transmitted infections in the USA.

In Black population centres in the USA, adult sex ratios (ASRs) are strongly female-biased primarily due to high male incarceration and early mortality rates. I explore the system of social determinants that shape these ASRs, and describe their apparent consequences. Evidence suggests that female-biased ASRs play a role, along with racial residential segregation, to increase mixing between core and peripheral members of sexual networks, facilitating transmission of human immunodeficiency virus and other sexually transmitted infections. Unique historical factors underlie Black male incarceration and mortality rates in the USA, making comparisons with other groups or other countries challenging.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'.

Persistent effect of sex ratios on relationship quality and life satisfaction.

Convict transportation to Australia imposed heavily male-biased sex ratios in some areas, altering the convict-era mating market and generating long-running cultural effects that persist to the present day. We test whether convict-era sex ratios have altered marital and overall life satisfaction today, through their persistent effects on gender norms and household bargaining. We find that both women and men are happier, and the happiness gap within married couples is smaller in areas where convict-era sex ratios were heavily male-biased than in areas where sex ratios were historically more even. We discuss our results in light of household bargaining theory, evolutionary sexual conflict theory and the well-documented relationship between conservative attitudes and self-reported happiness.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'.

Reproductive competition and sexual selection.

This paper traces the development of our understanding of the development of different approaches to estimating the strength of reproductive competition and sexual selection in the two sexes, based on measures of the operational sex ratio, the opportunity for sexual selection and contrasts in selection gradients between the sexes. It argues that different approaches provide complementary insights into the causes of sex differences in reproductive competition, the operation of sexual selection and the evolution of secondary sexual characters and that improvements in our understanding of the evolution of secondary sexual characters will require a more comprehensive understanding of the ways in which social and ecological conditions modify reproductive competition and development in females and males.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'.

High sex ratios in rural China: declining well-being with age in never-married men.

In parts of rural China male-biased sex ratios at birth, combined with out-migration of women, have led to highly male-biased adult sex ratios, resulting in large numbers of men being unable to marry, in a culture where marriage and reproduction are an expectation. The aim of this study was to test the hypotheses that older unmarried men are more predisposed to depression, low self-esteem and aggression than both those who are married, and those who are younger and unmarried. Self-completion questionnaires were administered among men aged 20–40 in 48 villages in rural Guizhou province, southwestern China. Tools used included the Beck Depression Inventory, the Rosenberg's Self-esteem Scale and the Bryant-Smith Aggression Questionnaire. Regression models assessed psychological wellbeing while adjusting for socio-demographic variables. Completed questionnaires were obtained from 957 never-married men, 535 married men aged 30–40, 394 partnered men and 382 unpartnered men aged 20–29. After adjusting for socio-demographic variables, never-married men were more predisposed to depression (p < 0.05), aggression (p < 0.01), low self-esteem (p < 0.05) and suicidal tendencies (p < 0.001). All the psychological measures deteriorated with age in never-married men. In contrast, married men remained stable on these dimensions with age. Never-married men are a psychologically highly vulnerable group in a society where marriage is an expectation. Since the highest birth sex–ratio cohorts have not yet reached reproductive age, the social tragedy of these men will last for at least another generation.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’.

Estimating adult sex ratios in nature.

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'.

Causes and consequences of adult sex ratio imbalance in a historical U.S. population.

The responsiveness of individuals to partner availability has been well-documented across the literature. However, there is disagreement regarding the direction of the consequences of sex ratio imbalance. Specifically, does an excess of males or females promote male-male mating competition? In an attempt to clarify the role of the adult sex ratio (ASR) on behaviour, here we evaluate both competing and complimentary expectations derived from theory across the social and biological sciences. We use data drawn from a historical, nineteenth century population in North America and target several life-history traits thought to be affected by partner availability: age at first birth, relationship status, completed fertility and longevity. Furthermore, we assess the role of various contributors to a population's ASR. We find that both the contributors to and consequences of sex ratio imbalance vary over time. Our results largely support predictions of greater male pairbond commitment and lesser male mating effort, as well as elevated bargaining power of women in response to female scarcity. After reviewing our findings, and others from across the literature, we highlight the need to adjust predictions in response to ASR imbalance by the: (i) culturally mediated mating arena, (ii) variable role of demographic inputs across time and place, (iii) constraints to behavioural outcomes across populations, and (iv) ability and accuracy of individuals to assess partner availability.This article is part of the themed issue 'Adult sex ratios and reproductive strategies: a critical re-examination of sex differences in human and animal societies'.

Adult sex ratio and social status predict mating and parenting strategies in Northern Ireland.

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.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'.

Not all sex ratios are equal: the Fisher condition, parental care and sexual selection.

The term 'sex roles' encapsulates male-female differences in mate searching, competitive traits that increase mating/fertilization opportunities, choosiness about mates and parental care. Theoretical models suggest that biased sex ratios drive the evolution of sex roles. To model sex role evolution, it is essential to note that in most sexually reproducing species (haplodiploid insects are an exception), each offspring has one father and one mother. Consequently, the total number of offspring produced by each sex is identical, so the mean number of offspring produced by individuals of each sex depends on the sex ratio (Fisher condition). Similarly, the total number of heterosexual matings is identical for each sex. On average, neither sex can mate nor breed more often when the sex ratio is even. But equally common in which sex ratio? The Fisher condition only applies to some reproductive measures (e.g. lifetime offspring production or matings) for certain sex ratios (e.g. operational or adult sex ratio; OSR, ASR). Here, we review recent models that clarify whether a biased OSR, ASR or sex ratio at maturation (MSR) have a causal or correlational relationship with the evolution of sex differences in parental care and competitive traits-two key components of sex roles. We suggest that it is more fruitful to understand the combined effect of the MSR and mortality rates while caring and competing than that of the ASR itself. In short, we argue that the ASR does not have a causal role in the evolution of parental care. We point out, however, that the ASR can be a cue for adaptive phenotypic plasticity in how each sex invests in parental care.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'.

Climate-driven shifts in adult sex ratios via sex reversals: the type of sex determination matters.

Sex reversals whereby individuals of one genetic sex develop the phenotype of the opposite sex occur in ectothermic vertebrates with 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.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'.

Adult sex ratios and their implications for cooperative breeding in birds.

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.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'.

Demographic and genetic consequences of disturbed sex determination

During sex determination, genetic and/or environmental factors determine the cascade of processes of gonad development. Many organisms, therefore, have a developmental window in which their sex determination can be sensitive to, for example, unusual temperatures or chemical pollutants. Disturbed environments can distort population sex ratios and may even cause sex reversal in species with genetic sex determination. The resulting genotype-phenotype mismatches can have long-lasting effects on population demography and genetics. I review the theoretical and empirical work in this context and explore in a simple population model the role of the fitness vyy of chromosomally aberrant YY genotypes that are a consequence of environmentally induced feminization. Low vyy is mostly beneficial for population growth. During feminization, low vyy reduces the proportion of genetic males and hence accelerates population growth, especially at low rates of feminization and at high fitness costs of the feminization itself (i.e. when feminization would otherwise not affect population dynamics much). When sex reversal ceases, low vyy mitigates the negative effects of feminization and can even prevent population extinction. Little is known about vyy in natural populations. The available models now need to be parametrized in order to better predict the long-term consequences of disturbed sex determination.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'.