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Mounting evidence suggests that patterns of local relatedness can change over time in predictable ways, a process termed kinship dynamics. Kinship dynamics may occur at the level of the population or social group, where the mean relatedness across all members of the population or group changes over time, or at the level of the individual, where an individual's relatedness to its local group changes with age. Kinship dynamics are likely to have fundamental consequences for the evolution of social behaviour and life history because they alter the inclusive fitness payoffs to actions taken at different points in time. For instance, growing evidence suggests that individual kinship dynamics have shaped the evolution of menopause and age-specific patterns of helping and harming. To date, however, the consequences of kinship dynamics for social evolution have not been widely explored. Here we review the patterns of kinship dynamics that can occur in natural populations and highlight how taking a kinship dynamics approach has yielded new insights into behaviour and life-history evolution. We discuss areas where analysing kinship dynamics could provide new insight into social evolution, and we outline some of the challenges in predicting and quantifying kinship dynamics in natural populations.

Rawls argued that fairness in human societies can be achieved if decisions about the distribution of societal rewards are made from behind a veil of ignorance, which obscures the personal gains that result. Whether ignorance promotes fairness in animal societies, that is, the distribution of resources to reduce inequality, is unknown. Here we show experimentally that cooperatively breeding banded mongooses, acting from behind a veil of ignorance over kinship, allocate postnatal care in a way that reduces inequality among offspring, in the manner predicted by a Rawlsian model of cooperation. In this society synchronized reproduction leaves adults in a group ignorant of the individual parentage of their communal young. We provisioned half of the mothers in each mongoose group during pregnancy, leaving the other half as matched controls, thus increasing inequality among mothers and increasing the amount of variation in offspring birth weight in communal litters. After birth, fed mothers provided extra care to the offspring of unfed mothers, not their own young, which levelled up initial size inequalities among the offspring and equalized their survival to adulthood. Our findings suggest that a classic idea of moral philosophy also applies to the evolution of cooperation in biological systems., Obscuring knowledge of personal gains from individuals can theoretically maintain fairness in a cooperative group. Experiments show that wild, cooperatively breeding banded mongooses uncertain of kinship allocate postnatal care in a way that reduces inequality among offspring, suggesting a classic idea of moral philosophy can apply in biological systems.

Most predictions on the evolution of adaptive parental effects and phenotypic memory exclusively focus on the role of the abiotic environment. How parental effects are affected by population demography and life history is less well understood. To overcome this, we use an analytical model to assess whether selection acting on fecundity versus viability affects the evolution of parental effects in a viscous population experiencing a spatiotemporally varying environment. We find that parental effects commonly evolve in regimes of viability selection, but are less likely to evolve in regimes of fecundity selection. In regimes of viability selection, an individual’s phenotype becomes correlated with its local environment during its lifetime, as those individuals with a locally adapted phenotype are more likely to survive until parenthood. Hence, a parental phenotype rapidly becomes an informative cue about its local environment, favoring the evolution of parental effects. By contrast, in regimes of fecundity selection, locally maladapted and adapted parents survive at equal rates, so that the parental phenotype, by itself, is not informative about the local environment. Correlations between phenotype and environment still arise, but only when more fecund, locally adapted individuals leave more successfully established offspring to the local patch. Hence, correlations take at least two generations to develop, making them more sensitive to distortion by environmental change or competition with immigrant offspring. Hence, we conclude that viability selection is most conducive to the evolution of adaptive parental effects in spatially structured populations.

Social groups whose members have had sustained prior experience with each other frequently exhibit improved coordination and outperform groups whose members are unfamiliar with one another. The mechanisms by which familiarity assists coordination are not well known. Prior social experience may simply allow individuals to learn the behavioral tendencies of familiar group-mates and coordinate accordingly. In the absence of prior social experience, it would be adaptive for individuals to develop strategies for coping with unfamiliar others to minimize the disadvantage of unfamiliarity. To explore the dynamics of familiarity in shaping group behaviors, we used a highly social catfish, Corydoras aeneus, that utilizes a distinctive, observable tactile interactions. Here we describe this tactile interaction behavior, physical “nudges” that are deployed to initiate group movements and maintain contact with group-mates during group movements. We then report the results of two experiments exploring the relationship between nudges and coordination. First, within triplets of two familiar and one unfamiliar individual, we found no individual differences in nudging rate based on familiarity. Despite all individuals interacting at similar rates, however, unfamiliar individuals failed to coordinate as well as their familiar group-mates, and were more frequently absent from group movements. Second, comparing pairs of familiar with pairs of unfamiliar fish, there was no difference in the level of coordination between pairs. Instead, we found that unfamiliar pairs exhibited significantly higher nudging rates, suggesting that unfamiliar pairs could compensate for their unfamiliarity by nudging more frequently. In contrast, familiar individuals coordinated with comparatively little nudging, presumably because they were experienced with each other. Overall, these results suggest that nudges can be used to improve coordination of group activities, but that their usage is reduced in the case of familiar individuals, implying that these potential signals may be costly.

Among species with variable numbers of individuals contributing to offspring care, an individual's investment strategy should depend upon both the size of the breeding group and the relative contributions of each carer. Existing theoretical work on carer investment rules has, however, largely focused on biparental care, and on modelling offspring provisioning in isolation from other stages of investment. Consequently, there has been little exploration of how maternal investment prior to birth might be expected to influence carer provisioning decisions after birth, and how these should be modified by the number of carers present. In particular, it is unclear whether mothers should increase or decrease their investment in each offspring under favourable rearing conditions, and whether this differs under alternative assumptions about the consequences of being ‘high quality’ at birth. We develop a game-theoretical model of cooperative care that incorporates female control of prebirth investment, and allow increased maternal investment to either substitute for later investment (giving offspring a ‘head start’) or raise the value of later investment (a ‘silver spoon’). We show that mothers reduce prebirth investment under better rearing conditions (more helpers) when investment is substitutable, leading to concealed helper effects. In contrast, when maternal prebirth investment primes offspring to benefit more from postbirth care, mothers should take advantage of good care environments by investing more in offspring both before and after birth. These results provide novel mechanisms to explain contrasting patterns of maternal investment across cooperative breeders.

Summary Why females of some species cease ovulation prior to the end of their natural lifespan is a long-standing evolutionary puzzle [1–4]. The fitness benefits of post-reproductive helping could in principle select for menopause [1, 2, 5], but the magnitude of these benefits appears insufficient to explain the timing of menopause [6–8]. Recent theory suggests that the cost of inter-generational reproductive conflict between younger and older females of the same social unit is a critical missing term in classical inclusive fitness calculations (the "reproductive conflict hypothesis" [6, 9]). Using a unique long-term dataset on wild resident killer whales, where females can live decades after their final parturition, we provide the first test of this hypothesis in a non-human animal. First, we confirm previous theoretical predictions that local relatedness increases with female age up to the end of reproduction. Second, we construct a new evolutionary model and show that given these kinship dynamics, selection will favor younger females that invest more in competition, and thus have greater reproductive success, than older females (their mothers) when breeding at the same time. Third, we test this prediction using 43 years of individual-based demographic data in resident killer whales and show that when mothers and daughters co-breed, the mortality hazard of calves from older-generation females is 1.7 times that of calves from younger-generation females. Intergenerational conflict combined with the known benefits conveyed to kin by post-reproductive females can explain why killer whales have evolved the longest post-reproductive lifespan of all non-human animals.

The handicap principle of Zahavi is potentially of great importance to the study of biological communication. Existing models of the handicap principle, however, make the unrealistic assumption that communication is error free. It seems possible, therefore, that Zahavi’s arguments do not apply to real signalling systems, in which some degree of error is inevitable. Here, we present a general evolutionarily stable strategy (ESS) model of the handicap principle which incorporates perceptual error. We show that, for a wide range of error functions, error-prone signalling systems must be honest at equilibrium. Perceptual error is thus unlikely to threaten the validity of the handicap principle. Our model represents a step towards greater realism, and also opens up new possibilities for biological signalling theory. Concurrent displays, direct perception of quality, and the evolution of ‘amplifiers9 and ‘attenuators’ are all probable features of real signalling systems, yet handicap models based on the assumption of error-free communication cannot accommodate these possibilities.

Evolutionarily stable strategy (ESS) models of biological signalling are important because the intimate coevolution of signalling and receiving strategies is complicated. Tentative results from a numerical study of error-prone signalling show the value of formal modelling. Error in perception can create discreteness in the distribution of signals produced, and so observed discreteness in nature may call for no more complicated explanation. Further developments in the theory of signalling may include a link with theories of aggression such as the sequential assessment game. The technical device of a ‘scratch space’ may allow a natural development of ‘two-way’ information games in which each contestant plays the roles of signaller and receiver simultaneously. This device may also incidentally derive mental states from purely strategic considerations.

In 1989, Hasson introduced the concept of an ‘amplifier’ within animal communication. This display reduces errors in the assessment of traits for which there is direct selection and renders differences in quality among animals more obvious. Amplifiers can evolve to fixation via the benefit they confer on high-quality animals. However, they also impose a cost on low-quality animals by revealing their lower quality, potentially leading these to refrain from amplifying. Hence, it was suggested that, if the level of amplification correlates with quality, direct choice for the amplifying display might emerge. Using the framework of signal detection theory, this article shows that, if the use of an amplifier is observable, direct choice for the amplifying display can indeed evolve. Consequently, low-quality animals may choose to amplify to some extent as well, even though this reveals their lower quality. In effect, the amplifier evolves to become a signal in its own right. We show that, as amplifiers can evolve without direct female choice and are likely to become correlated with male quality, selection for quality-dependent amplification provides a simple explanation for the origin of reliable signals in the absence of pre-existing preferences.

During collective movement, bolder individuals often emerge as leaders. Here, we investigate whether this reflects a greater propensity of bold individuals to initiate movement, or a preference for shy individuals to follow a bolder leader. We set up trios of stickleback fish comprising a focal individual who was either bold or shy, and one other individual of each personality. We then recorded the movements of all individuals in and out of cover in a foraging context to determine how assiduously the focal fish followed the movements of each other partner. We found that a shy focal fish preferred to follow a leader whose personality matched its own, but we did not detect such a difference in bold fish. Despite this preference, however, the greater propensity of bold individuals to initiate movements out of cover meant that they successfully led more joint trips. Thus, when offered a choice of leaders, sticklebacks prefer to follow individuals whose personality matches their own, but bolder individuals may, nevertheless, be able to impose their leadership, even among shy followers, simply through greater effort.

In this paper, we draw the attention of biologists to a result from the economic literature, which suggests that when individuals are engaged in a communal activity of benefit to all, selection may favour cooperative sharing of resources even among non-relatives. Provided that group members all invest some resources in the public good, they should refrain from conflict over the division of these resources. The reason is that, given diminishing returns on investment in public and private goods, claiming (or ceding) a greater share of total resources only leads to the actor (or its competitors) investing more in the public good, such that the marginal costs and benefits of investment remain in balance. This cancels out any individual benefits of resource competition. We illustrate how this idea may be applied in the context of biparental care, using a sequential game in which parents first compete with one another over resources, and then choose how to allocate the resources they each obtain to care of their joint young (public good) versus their own survival and future reproductive success (private good). We show that when the two parents both invest in care to some extent, they should refrain from any conflict over the division of resources. The same effect can also support asymmetric outcomes in which one parent competes for resources and invests in care, whereas the other does not invest but refrains from competition. The fact that the caring parent gains higher fitness pay-offs at these equilibria suggests that abandoning a partner is not always to the latter's detriment, when the potential for resource competition is taken into account, but may instead be of benefit to the ‘abandoned’ mate.

Human females stop reproducing long before they die. Among other mammals, only pilot and killer whales exhibit a comparable period of post-reproductive life. The grandmother hypothesis suggests that kin selection can favour post-reproductive survival when older females help their relatives to reproduce. But although there is an evidence that grandmothers can provide such assistance, it is puzzling why menopause should have evolved only among the great apes and toothed whales. We have previously suggested ( Cant & Johnstone 2008 Proc. Natl Acad. Sci. USA 105 , 5332–5336 ( doi:10.1073/pnas.0711911105 )) that relatedness asymmetries owing to female-biased dispersal in ancestral humans would have favoured younger females in reproductive competition with older females, predisposing our species to the evolution of menopause. But this argument appears inapplicable to menopausal cetaceans, which exhibit philopatry of both sexes combined with extra-group mating. Here, we derive general formulae for ‘kinship dynamics’, the age-related changes in local relatedness that occur in long-lived social organisms as a consequence of dispersal and mortality. We show that the very different social structures of great apes and menopausal whales both give rise to an increase in local relatedness with female age, favouring late-life helping. Our analysis can therefore help to explain why, of all long-lived, social mammals, it is specifically among the great apes and toothed whales that menopause and post-reproductive helping have evolved.

Low-level social aggression is a conspicuous feature of cooperative animal societies, but its precise function is usually unclear. One long-standing hypothesis is that aggressive displays by dominant individuals serve to reduce uncertainty about relative strength and deter subordinates from starting fights that they are unlikely to win. However, most formal theoretical models of this idea do not consider how the credibility of deterrent signals might change over time in social groups. We developed a simple model of dominant aggression as a deterrent signal, which takes into account how credibility changes over time and how selection should act on receiver memory. We then carried out an experimental test of the predictions of our model on a field population of the paper wasp, Polistes dominulus. The match between our theoretical and empirical results suggests that low-level social aggression can help to maintain the stability and productivity of cooperative associations in this species. Moreover, our work suggests that rates of aggression in animal societies and the robustness of social memories are likely to be intimately related.

In many animal groups, coordinated activity is facilitated by the emergence of leaders and followers. Although the identity of leaders is to some extent predictable, most groups experience frequent changes of leadership. How do group members cope with such changes in their social role? Here, we compared the foraging behaviour of pairs of stickleback fish after a period of either (i) role reinforcement , which involved rewarding the shyer follower for following, and the bolder leader for leading, or (ii) role reversal, which involved rewarding the shyer follower for leading, and the bolder leader for following. We found that, irrespective of an individual's temperament, its tendency to follow is malleable, whereas the tendency to initiate collective movement is much more resistant to change. As a consequence of this lack of flexibility in initiative, greater temperamental differences within a pair led to improved performance when typical roles were reinforced, but to impaired performance when typical roles were reversed.

In any system where multiple individuals jointly contribute to rearing offspring, conflict is expected to arise over the relative contributions of each carer. Existing theoretical work on the conflict over care has: (a) rarely considered the influence of tactical investment during offspring production on later contributions to offspring rearing; (b) concentrated mainly on biparental care, rather than cooperatively caring groups comprising both parents and helpers; and (c) typically ignored relatedness between carers as a potential influence on investment behavior. We use a game-theoretical approach to explore the effects of female production tactics and differing group relatedness structures on the expected rearing investment contributed by breeding females, breeding males, and helpers in cooperative groups. Our results suggest that the breeding female should pay higher costs overall when helpful helpers are present, as she produces additional offspring to take advantage of the available care. We find that helpers related to offspring through the breeding female rather than the breeding male should contribute less to care, and decrease their contribution as group size increases, because the female refrains from producing additional offspring to exploit them. Finally, within-group variation in helper relatedness also affects individual helper investment rules by inflating the differences between the contributions to care of dissimilar helpers. Our findings underline the importance of considering maternal investment decisions during offspring production to understand investment across the entire breeding attempt, and provide empirically testable predictions concerning the interplay between maternal, paternal and helper investment and how these are modified by different relatedness structures.

In cooperative breeding systems, some individuals help to raise offspring that are not their own. While early explanations for such altruistic behaviour were predominantly based on kin selection, recent evidence suggests that direct benefits may be important in the maintenance of cooperation. To date, however, discussions of cooperative breeding have made little reference to more general theories of cooperation between unrelated individuals (while these theories rarely address cooperative breeding). Here, we attempt to integrate the two fields. We identify four key questions that can be used to categorise different mechanisms for the maintenance of cooperative behaviour: (1) whether or not individuals invest in others; (2) whether or not this initial investment elicits a return investment by the beneficiary; (3) whether the interaction is direct, i.e. between two partners, or indirect (involving third parties) and (4) whether only actions that increase the fitness of the partner or also fitness reducing actions (punishment) are involved in the interaction. Asking these questions with regards to concepts in the literature on cooperative breeding, we found that (a) it is often straightforward to relate these concepts to general mechanisms of cooperation, but that (b) a single term (such as 'pay-to-stay', 'group augmentation' or 'prestige') may sometimes subsume two or more distinct mechanisms, and that (c) at least some mechanisms that are thought to be important in cooperative breeding systems have remained largely unexplored in the theoretical literature on the evolution of cooperation. Future theoretical models should incorporate asymmetries in power and pay off structure caused for instance by dominance hierarchies or partner choice, and the use of N-player games. The key challenges for both theoreticians and empiricists will be to integrate the hitherto disparate fields and to disentangle the parallel effects of kin and non-kin based mechanisms of cooperation.

Studies on leadership have focused either on physiological state as the key predictor (i.e. ‘‘leading according to need’’), or else on temperamental asymmetries among group members (i.e. intrinsic leadership). In this paper, we explore how both factors interact in determining the emergence of leaders. We observed pairs of sticklebacks with varying degrees of temperamental difference, and recorded their movements back and forth between a safe covered area and a risky foraging area, both before and after satiating one of the two pair members (but not the other). Before satiation, when the fish had similar hunger levels, temperament was a good predictor of social roles, with the bolder member of a pair leading and the shyer member following. The effect of satiation depended on which fish received the additional food. When the shyer member of a pair was fed, and consequently became less active, the bolder fish did not change its behaviour but continued to lead. By contrast, when the bolder member of a pair was fed, and consequently initiated fewer trips out of cover, the shyer partner compensated by initiating trips more frequently itself. In pairs that differed only a little in temperament, feeding the bolder fish actually led to a role reversal, with the shyer fish emerging as a leader in the majority of joint trips out of cover. Our results show that leadership emerges as the consequence of multiple factors, and that their interaction can be complex.

Current models of parent–offspring communication do not explicitly predict the effect of parental food supply on offspring demand (ESD). However, existing theory is frequently interpreted as predicting a negative ESD, such that offspring beg less when parental supply is high. While empirical evidence largely supports this interpretation, several studies have identified the opposite case, with well-fed offspring begging more than those in poorer condition. Here, we show that signalling theory can give rise to either a negative or a positive ESD depending on the precise form of costs and benefits. Introducing variation among parents in the cost of care, we show that the ESD may change sign depending upon the quantitative relation between two effects: (i) decreased supply leads to increased begging because of an increase in marginal fitness benefit of additional resources to offspring, (ii) decreased supply leads to reduced begging because it is associated with a decrease in parental responsiveness, rendering begging less effective. To illustrate the interplay between these two effects, we show that Godfray's seminal model of begging yields a negative ESD when care is generally cheap, because the impact of supply on the marginal benefits of additional resources then outweighs the associated changes in parental responsiveness to begging. By contrast, the same model predicts a positive ESD when care is generally costly, because the impact of care costs on parental responsiveness then outweighs the change in marginal benefits.

International audience; Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. Nowak et al. argue that inclusive fitness theory has been of little value in explaining the natural world, and that it has led to negligible progress in explaining the evolution of eusociality. However, we believe that their arguments are based upon a misunderstanding of evolutionary theory and a misrepresentation of the empirical literature. We will focus our comments on three general issues.

An enduring puzzle of human life history is why women cease reproduction midway through life. Selection can favor postreproductive survival because older females can help their offspring to reproduce. But the kin-selected fitness gains of helping appear insufficient to outweigh the potential benefits of continued reproduction. Why then do women cease reproduction in the first place? Here, we suggest that early reproductive cessation in humans is the outcome of reproductive competition between generations, and we present a simple candidate model of how this competition will be resolved. We show that among primates exhibiting a postreproductive life span, humans exhibit an extraordinarily low degree of reproductive overlap between generations. The rapid senescence of the human female reproductive system coincides with the age at which, in natural fertility populations, women are expected to encounter reproductive competition from breeding females of the next generation. Several lines of evidence suggest that in ancestral hominids, this younger generation typically comprised immigrant females. In these circumstances, relatedness asymmetries within families are predicted to give younger females a decisive advantage in reproductive conflict with older females. A model incorporating both the costs of reproductive competition and the benefits of grandmothering can account for the timing of reproductive cessation in humans and so offers an improved understanding of the evolution of menopause.

Background Foraging in groups offers animals a number of advantages, such as increasing their likelihood of finding food or detecting and avoiding predators. In order for a group to remain together, there has to be some degree of coordination of behaviour and movement between its members (which may in some cases be initiated by a decision-making leader, and in other cases may emerge as an underlying property of the group). For example, behavioural synchronisation is a phenomenon where animals within a group initiate and then continue to conduct identical behaviours, and has been characterised for a wide range of species. We examine how a pair of animals should behave using a state-dependent approach, and ask what conditions are likely to lead to behavioural synchronisation occurring, and whether one of the individuals is more likely to act as a leader. Results The model we describe considers how the energetic gain, metabolic requirements and predation risks faced by the individuals affect measures of their energetic state and behaviour (such as the degree of behavioural synchronisation seen within the pair, and the value to an individual of knowing the energetic state of its colleague). We explore how predictable changes in these measures are in response to changes in physiological requirements and predation risk. We also consider how these measures should change when the members of the pair are not identical in their metabolic requirements or their susceptibility to predation. We find that many of the changes seen in these measures are complex, especially when asymmetries exist between the members of the pair. Conclusion Analyses are presented that demonstrate that, although these general patterns are robust, care needs to be taken when considering the effects of individual differences, as the relationship between individual differences and the resulting qualitative changes in behaviour may be complex. We discuss how these results are related to experimental observations, and how the model and its predictions could be extended. Background: Foraging in groups offers animals a number of advantages, such as increasing their likelihood of finding food or detecting and avoiding predators. In order for a group to remain together, there has to be some degree of coordination of behaviour and movement between its members (which may in some cases be initiated by a decision-making leader, and in other cases may emerge as an underlying property of the group). For example, behavioural synchronisation is a phenomenon where animals within a group initiate and then continue to conduct identical behaviours, and has been characterised for a wide range of species. We examine how a pair of animals should behave using a state-dependent approach, and ask what conditions are likely to lead to behavioural synchronisation occurring, and whether one of the individuals is more likely to act as a leader. Results: The model we describe considers how the energetic gain, metabolic requirements and predation risks faced by the individuals affect measures of their energetic state and behaviour (such as the degree of behavioural synchronisation seen within the pair, and the value to an individual of knowing the energetic state of its colleague). We explore how predictable changes in these measures are in response to changes in physiological requirements and predation risk. We also consider how these measures should change when the members of the pair are not identical in their metabolic requirements or their susceptibility to predation. We find that many of the changes seen in these measures are complex, especially when asymmetries exist between the members of the pair. Conclusion: Analyses are presented that demonstrate that, although these general patterns are robust, care needs to be taken when considering the effects of individual differences, as the relationship between individual differences and the resulting qualitative changes in behaviour may be complex. We discuss how these results are related to experimental observations, and how the model and its predictions could be extended.

How can cooperation persist in the face of a temptation to 'cheat'? Several recent papers have suggested that the answer may lie in indirect reciprocity. Altruistic individuals may benefit by eliciting altruism from observers, rather than (as in direct reciprocity) from the recipient of the aid they provide. Here, we point out that indirect reciprocity need not always favour cooperation; by contrast, it may support spiteful behaviour, which is costly for the both actor and recipient. Existing theory suggests spite is unlikely to persist, but we demonstrate that it may do so when spiteful individuals are less likely to incur aggression from observers (a negative form of indirect reciprocity).

Close interactions between mother and offspring are said to result in a coevolution of parental and offspring genotypes such that offspring are adapted in their solicitation behaviour to obtain maternal provisioning that maximizes their fitness. Few empirical studies have been conducted in this field and it remains unclear whether maternal provisioning and offspring weight gain are influenced by the same set of maternal and offspring phenotypic and genotypic factors. Using a cross-foster, split-litter design in mice, we found that overall maternal provisioning and offspring weight gain are significantly correlated but are affected by a different set of parameters, except for the effect of maternal bodyweight. While the level of maternal provisioning was influenced by both offspring and foster mother genotype, offspring weight gain was only affected by the number of males in the mixed litter. We suggest that this disparity may hint at the inefficiency of offspring solicitation behaviour or effects of sibling competition.

Hominids display marked body size dimorphism, suggestive of strong sexual selection, yet they lack significant sex differences in canine size that are commonly associated with intrasexual competition in primates. We resolve this paradox by examining sex differences in hominoid facial morphology. We show that chimpanzees, but not gorillas, exhibit clear sexual dimorphism in face width, over and above that expected based on sex differences in body size. We show that this facial dimorphism, expressed as an index, is negatively correlated with canine dimorphism among anthropoid primates. Our findings suggest that a lack of canine dimorphism in anthropoids is not owing to weak sexual selection, but rather is associated with strong sexual selection for broader face width. Enlarged cheek-bones are linked with attractiveness in humans, and we propose that the evolution of a broad face and loss of large canines in hominid males results from mate choice.

Fights between pairs of animals frequently take place within a wider social context. The displays exchanged during conflict, and the outcome of an encounter, are often detectable by individuals who are not immediately involved. In at least some species, such bystanders are known to eavesdrop on contests between others, and to modify their behavior toward the contestants in response to the observed interaction. Here, I extend Maynard Smith's well known model of animal aggression, the Hawk–Dove game, to incorporate the possibility of eavesdroppers. I show that some eavesdropping is favored whenever the cost of losing an escalated fight exceeds the value of the contested resource, and that its equilibrium frequency is greatest when costs are relatively high. Eavesdropping reduces the risk of escalated conflict relative to that expected by chance, given the level of aggression in the population. However, it also promotes increased aggression, because it enhances the value of victory. The net result is that escalated conflicts are predicted to occur more frequently when eavesdropping is possible.

The study of quorum-sensing bacteria has revealed a widespread mechanism of coordinating bacterial gene expression with cell density. By monitoring a constitutively produced signal molecule, individual bacteria can limit their expression of group-beneficial phenotypes to cell densities that guarantee an effective group outcome. In this paper, we attempt to move away from a commonly expressed view that these impressive feats of coordination are examples of multicellularity in prokaryotic populations. Here, we look more closely at the individual conflict underlying this cooperation, illustrating that, even under significant levels of genetic conflict, signalling and resultant cooperative behaviour can stably exist. A predictive two-trait model of signal strength and of the extent of cooperation is developed as a function of relatedness (reflecting multiplicity of infection) and basic population demographic parameters. The model predicts that the strength of quorum signalling will increase as conflict (multiplicity of infecting strains) increases, as individuals attempt to coax more cooperative contributions from their competitors, leading to a devaluation of the signal as an indicator of density. Conversely, as genetic conflict increases, the model predicts that the threshold density for cooperation will increase and the subsequent strength of group cooperation will be depressed.

Previously developed models of reproductive skew have overlooked one of the main reasons why subordinates might remain in a group despite restricted opportunities to breed: the possibility of social queuing, i.e. acquiring dominant status in the future. Here, we present a dynamic ESS model of skew in animal societies that incorporates both immediate and future fitness consequences of the decisions taken by group members, based on their probability of surviving from one season to the next (when post-breeding survival probabilities drop to zero, our analysis reduces to the model produced by Reeve and Ratnieks in 1993, which considered only a single breeding season). This allows us to compare the delayed benefits of philopatry and the immediate opportunities for independent breeding. We show that delayed benefits greatly reduce the need for dominants to offer reproductive concessions to retain subordinates peacefully in the group. Moreover, this effect is strong enough that differences in survival have a much greater impact on the group structure than differences in other parameters, such as relatedness. When the possibility of acceding to dominant status is taken into account, groups where the dominant completely monopolizes reproduction can be stable, even if they consist of unrelated individuals, and even if subordinates have a reasonably high probability of winning a fight for dominance. Finally, we show that stable groups are possible even if association leads to a decrease in current productivity. Subordinates may still stand to gain from group membership under these circumstances, as acquiring breeding positions by queuing may be more efficient than the attempt to establish a new territory. At the same time, the dominant may be unable to exclude unwelcome subordinates, may enjoy increased survival when they are present, or may gain indirect benefits from allowing relatives to stay and queue for dominance. We conclude that reproductive skew in animal groups, ranging from eusocial insect colonies to mating aggregations (leks), will be strongly influenced by the future prospects of group members.

SummaryWhen individuals stand to gain by interacting with one another, but disagree over their preferred course of collective action, coordination can be hard to achieve [1–4]. In previous work, we found that pairs of stickleback fish prefer to synchronize their trips out of cover to look for food [5], possibly because this reduces perceived predation risk [6]. To create a degree of conflict over group coordination, we trained individual fish to expect food at one of two alternative, exposed locations and paired individuals with different expectations. Compared with isolated individuals, members of a pair showed a significantly increased tendency to alternate between foraging sites, together taking turns to visit first one individual's favored site and then the other individual's. Using a Markov-chain model to infer the individual rules underlying their joint behavior, we found that fish respond to a partner that breaks the pattern of alternation by themselves reverting to less regular behavior. Our results confirm theoretical predictions that conflict over group coordination can be resolved by taking turns [7–10] and show that, in this system, the pattern of alternation is actively monitored and maintained.