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

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Abstract

<p class="first" id="d6478943e169">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. </p><p id="d6478943e171">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’. </p>

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Natural Selection of Parental Ability to Vary the Sex Ratio of Offspring

R L Trivers, D. Willard (1973)

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Parental investment, sexual selection and sex ratios.

Hanna Kokko, Michael D. Jennions (2008)

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

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Sexual selection in males and females.

Tim H. Clutton-Brock (2007)

Research on sexual selection shows that the evolution of secondary sexual characters in males and the distribution of sex differences are more complex than was initially suggested but does not undermine our understanding of the evolutionary mechanisms involved. However, the operation of sexual selection in females has still received relatively little attention. Recent studies show that both intrasexual competition between females and male choice of mating partners are common, leading to strong sexual selection in females and, in extreme cases, to reversals in the usual pattern of sex differences in behavior and morphology.