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The transition from solitary life to sociality is considered one of the major transitions in evolution. In primates, this transition is currently not well understood. Traditional verbal models appear insufficient to unravel the complex interplay of environmental and demographic factors involved in the evolution of primate sociality, and recent phylogenetic reconstructions have produced conflicting results. We therefore analyze a theoretical model for the evolution of female social philopatry that sheds new light on the question why most primates live in groups. In individual-based simulations, we study the evolution of dispersal strategies of both resident females and their offspring. The model reveals that social philopatry can evolve through kin selection, even if retention of offspring is costly in terms of within-group resource competition and provides no direct benefits. Our model supports the role of predator avoidance as a selective pressure for group-living in primates, but it also suggests that a second benefit of group-living, communal resource defense, might be required to trigger the evolution of sizable groups. Lastly, our model reveals that seemingly small differences in demographic parameters can have profound effects on primate social evolution., (© 2020 The Authors. American Journal of Physical Anthropology published by Wiley Periodicals LLC.)

Theory suggests that genetic conflicts drive turnovers between sex-determining mechanisms, yet these studies only apply to cases where sex allocation is independent of environment or condition. Here, we model parent-offspring conflict in the presence of condition-dependent sex allocation, where the environment has sex-specific fitness consequences. Additionally, one sex is assumed to be more costly to produce than the other, which leads offspring to favor a sex ratio less biased toward the cheaper sex in comparison to the sex ratio favored by mothers. The scope for parent-offspring conflict depends on the relative frequency of both environments: when one environment is less common than the other, parent-offspring conflict can be reduced or even entirely absent, despite a biased population sex ratio. The model shows that conflict-driven invasions of condition-independent sex factors (e.g., sex chromosomes) result either in the loss of condition-dependent sex allocation, or, interestingly, lead to stable mixtures of condition-dependent and condition-independent sex factors. The latter outcome corresponds to empirical observations in which sex chromosomes are present in organisms with environment-dependent sex determination. Finally, conflict can also favor errors in environmental perception, potentially resulting in the loss of condition-dependent sex allocation without genetic changes to sex-determining loci., (© 2014 The Author(s). Evolution © 2014 The Society for the Study of Evolution.)

Sex allocation theory predicts the optimal allocation to male and female reproduction in sexual organisms. In animals, most work on sex allocation has focused on species with separate sexes and our understanding of simultaneous hermaphrodites is patchier. Recent theory predicts that sex allocation in simultaneous hermaphrodites should strongly be affected by post-copulatory sexual selection, while the role of pre-copulatory sexual selection is much less clear. Here, we review sex allocation and sexual selection theory for simultaneous hermaphrodites, and identify several strong and potentially unwarranted assumptions. We then present a model that treats allocation to sexually selected traits as components of sex allocation and explore patterns of allocation when some of these assumptions are relaxed. For example, when investment into a male sexually selected trait leads to skews in sperm competition, causing local sperm competition, this is expected to lead to a reduced allocation to sperm production. We conclude that understanding the evolution of sex allocation in simultaneous hermaphrodites requires detailed knowledge of the different sexual selection processes and their relative importance. However, little is currently known quantitatively about sexual selection in simultaneous hermaphrodites, about what the underlying traits are, and about what drives and constrains their evolution. Future work should therefore aim at quantifying sexual selection and identifying the underlying traits along the pre- to post-copulatory axis.

In most species, some individuals delay reproduction or occupy inferior breeding positions. The queue hypothesis tries to explain both patterns by proposing that individuals strategically delay breeding (queue) to acquire better breeding or social positions. In 1995, Ens, Weissing, and Drent addressed evolutionarily stable queuing strategies in situations with habitat heterogeneity. However, their model did not consider the non-mutually exclusive individual quality hypothesis, which suggests that some individuals delay breeding or occupy inferior breeding positions because they are poor competitors. Here we extend their model with individual differences in competitive abilities, which are probably plentiful in nature. We show that including even the smallest competitive asymmetries will result in individuals using queuing strategies completely different from those in models that assume equal competitors. Subsequently, we investigate how well our models can explain settlement patterns in the wild, using a long-term study on oystercatchers. This long-lived shorebird exhibits strong variation in age of first reproduction and territory quality. We show that only models that include competitive asymmetries can explain why oystercatchers' settlement patterns depend on natal origin. We conclude that predictions from queuing models are very sensitive to assumptions about competitive asymmetries, while detecting such differences in the wild is often problematic.

We investigate the conflict between queen and worker over sex allocation, specifically the allocation of the queen's eggs between workers and reproductives and the allocation of the reproductive eggs between male and female. In contrast to previous models, we allow workers to observe and use information about the strategy of the queen. We consider three conflict models: simultaneous (no information exchange), sequential (a one-way information exchange) and negotiated (an iterated two-way information exchange). We find that the first model produces sex ratios intermediate between the classic queen (1:1) and worker (1:3) optima. The second model, in which the worker has information about the queen's decisions, produces a different result and one that is somewhat counter-intuitive in that the sex ratios are less female-biased than for the other two models, and in fact are often male-biased. The third model predicts sex ratios intermediate between the first two models. We discuss how these findings may shed new light on observed sex allocation patterns in social insects and we suggest some experimental tests.

Sex ratio theory provides a clear and simple way to test if nonsocial haplodiploid wasps can discriminate between kin and nonkin. Specifically, if females can discriminate siblings from nonrelatives, then they are expected to produce a higher proportion of daughters if they mate with a sibling. This prediction arises because in haplodiploids, inbreeding (sib-mating) causes a mother to be relatively more related to her daughters than her sons. Here we formally model this prediction for when multiple females lay eggs in a patch, and test it with the parasitoid wasp Nasonia vitripennis. Our results show that females do not adjust their sex ratio behaviour dependent upon whether they mate with a sibling or nonrelative, in response to either direct genetic or a range of indirect environmental cues. This suggests that females of N. vitripennis cannot discriminate between kin and nonkin. The implications of our results for the understanding of sex ratio and social evolution are discussed.

We review some recent theoretical and empirical developments in the study of sex allocation in birds. The advent of reliable molecular sexing techniques has led to a sharp increase in the number of studies that report biased offspring sex ratios in birds. However, compelling evidence for adaptive sex allocation in birds is still very scant. We argue that there are two reasons for this: (i) standard sex allocation models, very helpful in understanding sex allocation of invertebrates, do not sufficiently take the complexities of bird life histories and physiology into account. Recent theoretical work might bring us a step closer to more realistic models; (ii) experimental field and laboratory studies on sex allocation in birds are scarce. Recent experimental work both in the laboratory and in the field shows that this is a promising approach.