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Collective conflicts among humans are widespread, although often highly destructive. A classic explanation for the prevalence of such warfare in some human societies is leadership by self-serving individuals that reap the benefits of conflict while other members of society pay the costs. Here, we show that leadership of this kind can also explain the evolution of collective violence in certain animal societies. We first extend the classic hawk-dove model of the evolution of animal aggression to consider cases in which a subset of individuals within each group may initiate fights in which all group members become involved. We show that leadership of this kind, when combined with inequalities in the payoffs of fighting, can lead to the evolution of severe intergroup aggression, with negative consequences for population mean fitness. We test our model using long-term data from wild banded mongooses, a species characterized by frequent intergroup conflicts that have very different fitness consequences for male and female group members. The data show that aggressive encounters between groups are initiated by females, who gain fitness benefits from mating with extragroup males in the midst of battle, whereas the costs of fighting are borne chiefly by males. In line with the model predictions, the result is unusually severe levels of intergroup violence. Our findings suggest that the decoupling of leaders from the costs that they incite amplifies the destructive nature of intergroup conflict.

The ultimate payoff of behaviours depends not only on their direct impact on an individual, but also on the impact on their relatives. Local relatedness-the average relatedness of an individual to their social environment-therefore has profound effects on social and life history evolution. Recent work has begun to show that local relatedness has the potential to change systematically over an individual's lifetime, a process called kinship dynamics. However, it is unclear how general these kinship dynamics are, whether they are predictable in real systems and their effects on behaviour and life history evolution. In this study, we combine modelling with data from real systems to explore the extent and impact of kinship dynamics. We use data from seven group-living mammals with diverse social and mating systems to demonstrate not only that kinship dynamics occur in animal systems, but also that the direction and magnitude of kinship dynamics can be accurately predicted using a simple model. We use a theoretical model to demonstrate that kinship dynamics can profoundly affect lifetime patterns of behaviour and can drive sex differences in helping and harming behaviour across the lifespan in social species. Taken together, this work demonstrates that kinship dynamics are likely to be a fundamental dimension of social evolution, especially when considering age-linked changes and sex differences in behaviour and life history.

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.

The extended female postreproductive life span found in humans and some toothed whales remains an evolutionary puzzle. Theory predicts demographic patterns resulting in increased female relatedness with age (kinship dynamics) can select for a prolonged postreproductive life span due to the combined costs of intergenerational reproductive conflict and benefits of late‐life helping. Here, we test this prediction using >40 years of longitudinal demographic data from the sympatric yet genetically distinct killer whale ecotypes: resident and Bigg's killer whales. The female relatedness with age is predicted to increase in both ecotypes, but with a less steep increase in Bigg's due to their different social structure. Here, we show that there is a significant postreproductive life span in both ecotypes with >30% of adult female years being lived as postreproductive, supporting the general prediction that an increase in local relatedness with age predisposes the evolution of a postreproductive life span. Differences in the magnitude of kinship dynamics however did not influence the timing or duration of the postreproductive life span with females in both ecotypes terminating reproduction before their mid‐40s followed by an expected postreproductive period of about 20 years. Our results highlight the important role of kinship dynamics in the evolution of a long postreproductive life span in long‐lived mammals, while further implying that the timing of menopause may be a robust trait that is persistent despite substantial variation in demographic patterns among populations., The rare life history trait of a long postreproductive period in females have been linked to demographic patterns leading to an increased relatedness to the local group with age (kinship dynamics). Here, we show that two killer whale ecotypes that are genetically distinct and exhibit different demographic patterns as well as different strengths of expected kinship dynamics have evolved a long postreproductive period in females, supporting the prediction that an increased female local relatedness with age predisposes for the evolution of this life history trait. Moreover, this results indicates that the onset of the postreproductive period may be robust to substantial variation in demographic patterns among 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.

Numerous studies have shown that social adversity in early life can have long-lasting consequences for social behaviour in adulthood, consequences that may in turn be propagated to future generations. Given these intergenerational effects, it is puzzling why natural selection might favour such sensitivity to an individual’s early social environment. To address this question, we model the evolution of social sensitivity in the development of helping behaviours, showing that natural selection indeed favours individuals whose tendency to help others is dependent on early-life social experience. In organisms with non-overlapping generations, we find that natural selection can favour positive social feedbacks, in which individuals who received more help in early life are also more likely to help others in adulthood, while individuals who received no early-life help develop low tendencies to help others later in life. This positive social sensitivity is favoured because of an intergenerational relatedness feedback: patches with many helpers tend to be more productive, leading to higher relatedness within the local group, which in turn favours higher levels of help in the next generation. In organisms with overlapping generations, this positive feedback is less likely to occur, and those who received more help may instead be less likely to help others (negative social feedback). We conclude that early-life social influences can lead to strong between-individual differences in helping behaviour, which can take different forms dependent on the life history in question. This article is part of the theme issue ‘Developing differences: early-life effects and evolutionary medicine’.

Johnstone and Cant introduce menopause and the phenomenon of an extended post-reproductive life found in humans and very few other mammals.

© 2018 by The University of Chicago. The relationships between mating decisions and parental investment are central to evolution, but to date few theoretical treatments of their coevolution have been developed. Here we adopt a demographically explicit, adaptive dynamics approach to analyze the coevolution of female mating decisions and parental investment of both sexes in a self-consistent way. Our models predict that where females cannot interfere with one another’s mating decisions and where they do not differ in their survival and fecundity prospects, monogamy should be rare, favored only under harsh environmental conditions, in sparse populations. However, allowing for interference or asymmetries among females leads to selection for monogamy over a much broader range of environments and demographies. Interference by paired, resident females may prevent unmated rivals from joining existing monogamous pairs, thus barring the formation of polygynous groups. Asymmetries between established, primary females and subsequently joining secondary females may increase the relative costs of early polygynous reproduction, compared to delayed monogamy for the latter. The models thus highlight different routes by which monogamy may evolve. We further track how parental investment by the sexes coevolves with female mating decisions, highlighting how sexual conflict over parental investment is both cause and effect of mating behavior.

Existing theory on the evolution of parental effects and the inheritance of non-genetic factors has mostly focused on the role of environmental change. By contrast, how differences in population demography and life history affect parental effects is poorly understood. To fill this gap, we develop an analytical model to explore how parental effects evolve when selection acts on fecundity versus viability in spatio-temporally fluctuating environments. We find that regimes of viability selection, but not fecundity selection, are most likely to favour parental effects. In the case of viability selection, locally adapted phenotypes have a higher survival than maladapted phenotypes and hence become enriched in the local environment. Hence, simply by being alive, a parental phenotype becomes correlated to its environment (and hence informative to offspring) during its lifetime, favouring the evolution of parental effects. By contrast, in regimes of fecundity selection, correlations between phenotype and environment develop more slowly: this is because locally adapted and maladapted parents survive at equal rates (no survival selection), so that parental phenotypes, by themselves, are uninformative about the local environment. However, because locally adapted parents are more fecund, they contribute more offspring to the local patch than maladapted parents. In case these offspring are also likely to inherit the adapted parents’ phenotypes (requiring pre-existing inheritance), locally adapted offspring become enriched in the local environment, resulting in a correlation between phenotype and environment, but only in the offspring’s generation. Because of this slower build-up of a correlation between phenotype and environment essential to parental effects, fecundity selection is more sensitive to any distortions owing to environmental change than viability selection. Hence, we conclude that viability selection is most conducive to the evolution of parental effects. This article is part of the theme issue ‘How does epigenetics influence the course of evolution?’

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.

Established mimicry theory predicts that Batesian mimics are selected to resemble their defended models, while models are selected to become dissimilar from their mimics. However, this theory has mainly considered individual selection acting on solitary organisms such as adult butterflies. Although Batesian mimicry of social insects is common, the few existing applications of kin selection theory to mimicry have emphasized relatedness among mimics rather than among models. Here, we present a signal detection model of Batesian mimicry in which the population of defended model prey is kin structured. Our analysis shows for most of parameter space that increased average dissimilarity from mimics has a twofold group-level cost for the model prey: it attracts more predators and these adopt more aggressive attack strategies. When mimetic resemblance and local relatedness are sufficiently high, such costs acting in the local neighbourhood may outweigh the individual benefits of dissimilarity, causing kin selection to drive the models to resemble their mimics. This requires model prey to be more common than mimics and/or well-defended, the conditions under which Batesian mimicry is thought most successful. Local relatedness makes defended prey easier targets for Batesian mimicry and is likely to stabilize the mimetic relationship over time.

Cooperative societies vary in the extent to which reproduction is skewed toward one or a few socially dominant animals. Many recent models attempt to explain this variation on the basis that a dominant who benefits from the presence of subordinates may offer them incentives, in the form of reproductive opportunities, to remain in the group. While most societies contain multiple members, however, these models have considered only the relationship between a dominant and a single subordinate or have assumed that all subordinates are identical. We develop an incentive-based evolutionary stable strategy model of reproductive skew in three-member groups, in which subordinates may vary in their opportunities for independent reproduction, their contribution to group productivity, and in their relatedness both to the dominant and to one another. Our model demonstrates that the conclusions of two-member models cannot all be generalized to larger groups. For example, relatedness among group members can influence whether or not the dominant does best to offer staying incentives to subordinates in a three-member, but not a two-member, group. Both the degree of skew and group stability depend on the relatedness between subordinates as well as on the relatedness of each to the dominant, and the incentives that each individual subordinate receives are influenced by the traits of the other. Whether such effects increase or decrease skew and group stability depends crucially on whether a third group member increases group productivity to a greater or lesser extent than the first.

Despite growing evidence for nongenetic inheritance, the ecological conditions that favor the evolution of heritable parental or grandparental effects remain poorly understood. Here, we systematically explore the evolution of parental effects in a patch-structured population with locally changing environments. When selection favors the production of a mix of offspring types, this mix differs according to the parental phenotype, implying that parental effects are favored over selection for bet-hedging in which the mixture of offspring phenotypes produced does not depend on the parental phenotype. Positive parental effects (generating a positive correlation between parental and offspring phenotype) are favored in relatively stable habitats and when different types of local environment are roughly equally abundant, and can give rise to long-term parental inheritance of phenotypes. By contrast, unstable habitats can favor negative parental effects (generating a negative correlation between parental and offspring phenotype), and under these circumstances even slight asymmetries in the abundance of local environmental states select for marked asymmetries in transmission fidelity.

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.

Maternal effects can provide offspring with reliable information about the environment they are likely to experience, but also offer scope for maternal manipulation of young when interests diverge between parents and offspring. To predict the impact of parent-offspring conflict, we model the evolution of maternal effects on local adaptation of young. We find that parent-offspring conflict strongly influences the stability of maternal effects; moreover, the nature of the disagreement between parents and young predicts how conflict is resolved: when mothers favor less extreme mixtures of phenotypes relative to offspring (i.e., when mothers stand to gain by hedging their bets), mothers win the conflict by providing offspring with limited amounts of information. When offspring favor overproduction of one and the same phenotype across all environments compared to mothers (e.g., when offspring favor a larger body size), neither side wins the conflict and signaling breaks down. Only when offspring favor less extreme mixtures relative to their mothers (something no current model predicts), offspring win the conflict and obtain full information about the environment. We conclude that a partial or complete breakdown of informative maternal effects will be the norm rather than the exception in the presence of parent-offspring conflict.

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.

For individuals collaborating to rear offspring, effective organization of resource delivery is difficult because each carer benefits when the others provide a greater share of the total investment required. When investment is provided in discrete events, one possible solution is to adopt a turn-taking strategy whereby each individual reduces its contribution rate after investing, only increasing its rate again once another carer contributes. To test whether turn-taking occurs in a natural cooperative care system, here we use a continuous time Markov model to deduce the provisioning behavior of the chestnut-crowned babbler (Pomatostomus ruficeps), a cooperatively breeding Australian bird with variable number of carers. Our analysis suggests that turn-taking occurs across a range of group sizes (2–6), with individual birds being more likely to visit following other individuals than to make repeat visits. We show using a randomization test that some of this apparent turn-taking arises as a by-product of the distribution of individual inter-visit intervals (“passive” turn-taking) but that individuals also respond actively to the investment of others over and above this effect (“active” turn-taking). We conclude that turn-taking in babblers is a consequence of both their individual provisioning behavior and deliberate response rules, with the former effect arising through a minimum interval required to forage and travel to and from the nest. Our results reinforce the importance of considering fine-scale investment dynamics when studying parental care and suggest that behavioral rules such as turn-taking may be more common than previously thought. Significance statement Caring for offspring is a crucial stage in the life histories of many animals and often involves conflict as each carer typically benefits when others contribute a greater share of the work required. One way to resolve this conflict is to monitor when other carers contribute and adopt a simple “turn-taking” rule to ensure fairness, but natural parental care has rarely been studied in sufficient detail to identify such rules. Our study investigates whether cooperatively breeding chestnut-crowned babblers “take turns” delivering food to offspring, and (if so) whether this a deliberate strategy or simply a by-product of independent care behavior. We find that babblers indeed take turns and conclude that part of the observed turn-taking is due to deliberate responsiveness, with the rest arising from the species’ breeding ecology. Electronic supplementary material The online version of this article (10.1007/s00265-017-2391-4) contains supplementary material, which is available to authorized users.

The emergence of leaders and followers is a key factor in facilitating group cohesion in animals. Individual group members have been shown to respond strongly to each other’s behavior and thereby affect the emergence and maintenance of these social roles. However, it is not known to what extent previous social experience might still affect individual’s leading and following tendencies in later social interactions. Here, by pairing three-spined sticklebacks (Gasterosteus aculeatus) with 2 different consecutive partners, we show a carryover effect of a previous partner’s personality on the behavior of focal individuals when paired with a new partner. This carryover effect depended on the relative boldness of the focal individual. Relatively bold but not shy fish spent less time out of cover and led their current partner less if they had previously been paired with a bolder partner. By contrast, following behavior was mainly influenced by the personality of the current partner. Overall, the behavior of relatively bold fish was more consistent across the stages, whereas shy fish changed their behavior more strongly depending on the current context. These findings emphasize how the history of previous social interactions can play a role in the emergence and maintenance of social roles within groups, providing an additional route for individual differences to affect collective behavior.

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.

When individuals invest in a common good, an efficient outcome is hard to achieve, because each can free ride on others’ efforts. This problem can lead parents that raise their young together to reduce their investment in care, with negative consequences for offspring. Here, we present a mathematical model to show that a strategy of conditional cooperation, in which parents take turns feeding their young, can help to resolve this problem. To test this idea, we studied the behavior of great tit parents raising chicks together. We found that parents alternated visits to the nest more than would be expected by chance, speeding up their feeding rate after their partner had visited the chicks, but slowing down again once they had visited in turn. This effect was not mediated by visit-to-visit changes in offspring begging intensity, although females (but not males) were influenced by mean begging levels across broods. We conclude that conflict over parental investment in this species is partly ameliorated by a simple form of reciprocity.

Social groups composed of familiar individuals exhibit better coordination than unfamiliar groups; however, the ways familiarity contributes to coordination are poorly understood. Prior social experience probably allows individuals to learn the tendencies of familiar group-mates and respond accordingly. Without prior experience, individuals would benefit from strategies for enhancing coordination with unfamiliar others. We used a social catfish, Corydoras aeneus , that uses discrete, observable tactile interactions to assess whether active interactions could facilitate coordination, and how their role might be mediated by familiarity. We describe this previously understudied physical interaction, ‘nudges’, and show it to be associated with group coordination and cohesion. Furthermore, we investigated nudging and coordination in familiar/unfamiliar pairs. In all pairs, we found that nudging rates were higher during coordinated movements than when fish were together but not coordinating. We observed no familiarity-based difference in coordination or cohesion. Instead, unfamiliar pairs exhibited significantly higher nudging rates, suggesting that unfamiliar pairs may be able to compensate for unfamiliarity through increased nudging. By contrast, familiar individuals coordinated with comparatively little nudging. Second, we analysed nudging and cohesion within triplets of two familiar and one unfamiliar individual (where familiar individuals had a choice of partner). Although all individuals nudged at similar rates, the unfamiliar group-mate was less cohesive than its familiar group-mates and spent more time alone. Unfamiliar individuals that nudged their group-mates more frequently exhibited higher cohesion, indicating that nudging may facilitate cohesion for the unfamiliar group-mate. Overall, our results suggest that nudges can mitigate unfamiliarity, but that their usage is reduced in the case of familiar individuals, implying a cost is associated with the behaviour.

SummaryThe study of animal signals began in earnest with the publication in 1872 of Charles Darwin’s The Expressions of the Emotions in Man and Animals, which laid the basis for a comparative study of signals across all animals, including humans. Yet even before Darwin, the exceptional diversity of animal signals has gripped the attention of natural historians and laymen alike, as these signals represent some of the most striking features of the natural world. Structures such as the long ornamented tail of the peacock, the roaring sounds of howler monkeys, audible kilometers away, and the pheromone trails laid by ants to guide their nestmates to resources are each examples of animal signals (Figure 1). Indeed, because signals evolved for the purpose of communicating (Box 1), their prominence can be hard for even a casual observer to overlook. Animal signals therefore raise many scientific questions: What are their functions? What information do they transmit? How are they produced? And why did they evolve?

Recent theory in social evolution has been mainly concerned with competition and cooperation within social groups of animals and their impact on the stability of those groups. Much less attention has been paid to conflicts arising as a result of solitary floaters (outsiders) attempting to join groups of established residents (insiders). We model such conflicts over group-membership using a demographically explicit approach in which the rates of births and deaths in a population determine the availability of group-vacancies and the number of floaters competing over these vacancies. We find that the outcome of within-group competition, reflected in the partitioning of reproduction among group members, exerts surprisingly little influence on the resolution of insider-outsider conflict. The outcome of such conflict is also largely unaffected by differences in resource holding potential between insiders and outsiders. By contrast, whether or not groups form is mainly determined by demographic factors (variation in vital rates such as fecundity and mortality) and the resulting population dynamics. In particular, at high floater densities territory defense becomes too costly, and groups form because insiders give in to the intruder pressure imposed on them by outsiders. We emphasize the importance of insider-outsider conflicts in social evolution theory and highlight avenues for future research.

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.

Theoretical advances in the study of social evolution have highlighted the importance of studying group formation in conjunction with population dynamics. To address this need, a number of demographically explicit models have been developed. We parameterize such a model to study the evolution of multimale associations in Verreaux’s sifaka (Propithecus verreauxi), a diurnal lemur living in small multimale–multifemale groups. We use long-term demographic data from a population in western Madagascar to estimate the fitness prospects of different male behavioral strategies. Our results indicate that dispersing male sifaka, rather than searching for a breeding territory of their own (floating), do better joining an established group as a subordinate with limited reproductive opportunities, mainly because they may later ascend to the dominant position. By contrast, resident males should not accept potential immigrants because we could not find any measurable benefit of living in a multimale group for them. There is thus a conflict over group membership between resident and floater males, and we suggest that floaters at least occasionally win this conflict because they have more to gain from joining than residents stand to lose by sharing their territory. Key words: ecological constraints, group-living, Propithecus verreauxi, social evolution, social queuing. [Behav Ecol]

Models that investigate family conflicts over parental investment have generally ignored population structure above the level of the family. Here, we extend conventional models to assess the impact of spatial population structure and limited dispersal on the intensity and resolution of parent‐offspring conflict. Although one might naively predict that limited dispersal should reduce parent‐offspring conflict, due to increased local relatedness, we show that this is not necessarily the case. Parent‐offspring conflict may actually become more intense when dispersal is limited because local competition among related young can reduce the cost of any decline in maternal productivity, favoring greater selfishness on the part of the offspring. We also investigate the impact of limited dispersal on the resolution of parent‐offspring conflict when the young can solicit extra resources by means of costly begging. Our model predicts that begging will be more intense when dispersal is limited but that the impact of dispersal on size then depends on the effect of parental supply on offspring demand. When greater supply leads to reduced demand, we find that parental investment should be higher in more viscous populations. By contrast, when greater supply leads to greater demand, parental investment should be lower in more viscous populations. Key words: begging, kin selection, local competition, model, offspring demand. [Behav Ecol]

In his famous haplodiploidy hypothesis, W. D. Hamilton proposed that high sister–sister relatedness facilitates the evolution of kin-selected reproductive altruism among Hymenopteran females. Subsequent analyses, however, suggested that haplodiploidy cannot promote altruism unless altruists capitalize on relatedness asymmetries by helping to raise offspring whose sex ratio is more female-biased than the population at large. Here, we show that haplodiploidy is in fact more favourable than is diploidy to the evolution of reproductive altruism on the part of females, provided only that dispersal is male-biased (no sex-ratio bias or active kin discrimination is required). The effect is strong, and applies to the evolution both of sterile female helpers and of helping among breeding females. Moreover, a review of existing data suggests that female philopatry and non-local mating are widespread among nest-building Hymenoptera. We thus conclude that Hamilton was correct in his claim that ‘family relationships in the Hymenoptera are potentially very favourable to the evolution of reproductive altruism’.

When members of a group differ in their preferred course of action, coordination poses a challenge. Leadership offers one way to resolve this difficulty, but the evolution of leaders and followers is itself poorly understood. Existing discussions have frequently attributed leadership to differences in information or need among individuals. Here, however, we show that in an n -player, repeated coordination game, selection leads to evolutionary branching and diversification in intrinsic leadership among the members of a population even in the absence of any variation in state. When individuals interact in pairs, repeated branching is possible; when individuals interact in larger groups, the typical outcome is a single branching event leading to a dimorphism featuring extreme intrinsic leaders and followers. These personality types emerge and are maintained by frequency-dependent selection, because leaders gain by imposing their preferences on followers, but fail to coordinate effectively when interacting with other leaders. The fraction of intrinsic leaders in the population increases with the degree of conflict among group members, with both types common only at intermediate levels of conflict; when conflict is weak, most individuals are intrinsic followers, and groups achieve high levels of coordination by randomly converging on one individual's preferred option, whereas when conflict is strong, most individuals are intrinsic leaders, and coordination breaks down because members of a group are rarely willing to follow another.

In cooperative breeders, parents that receive help with offspring care may either maintain their own effort or reduce it in compensation so that offspring gain little. Here, I build on an approach developed by McNamara et al. (McNamara, Gasson, and Houston 1999. Incorporating rules for responding into evolutionary games. Nature. 401: 368--371 and McNamara, Szekely, Webb, and Houston 2003. Should young ever be better off with one parent than with two? Behav Ecol. 14: 301--310.) to model behavioral negotiation over care in a cooperatively breeding species. I show that parental responsiveness to helper effort should vary with the shape of the parental cost function and that sex differences in the costs of care can give rise to sex differences in response to helpers. Such differences in responsiveness have major implications for the evolution of helping effort. When load lightening occurs, help may benefit mothers or fathers as much as offspring and to different degrees depending on the behavior of the 2. Consequently, when considering the influence of relatedness on helping effort, it is not only the relatedness of helpers to the young they assist that matters. Whether maternal or paternal kin help more, for instance, should reflect the relative degree of compensation by female and male parents. To understand helping, one must consider the whole pattern of relatedness among family members, as well as the ways in which they respond to one another. Copyright 2011, Oxford University Press.

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.

In foraging groups, individuals may utilise information from their social environment to aid decision making when choosing where to search for food. Little work has looked at the costs or benefits of behavioural differences, such as consistent individual variation in boldness, with respect to learning ability. Here, we investigate the response of three-spined stickleback (Gasterosteus aculeatus) to ‘social cues’, ‘local enhancement’ and ‘public information’ during foraging tasks. Our results confirm previous work suggesting that this species responds to social cues and local enhancement but not public information. Variation in boldness did not affect the use of different types of information. However, time taken to make a choice and reach a patch varied between fish with different levels of boldness. Contrary to expectation, shy fish were the more variable individuals, having a greater range of reaction times when responding to the tasks. This suggests that individual behavioural differences still play a role when utilising information obtained from the environment and may influence the relative benefits that could result in different contexts.

Speak to Me, MamaMaternal effects are effects of the mother (for example, hormones) on her offspring that are independent of genetic inheritance, but related to the mother's phenotype and her environment. Despite their universal occurrence, little is known about the evolutionary influence of maternal effects.Hindeet al.(p.1373; see the cover) modeled maternal effects in birds and showed that nestling begging, as well as provoking feeding, also provides parents with information about offspring vigor. Conversely, maternal effects provide information to the unborn offspring about parental quality. Experimental studies with canaries provided support for the theoretical predictions, and together these studies show that parent-offspring conflict (that is, the conflict between the offspring for its immediate needs and the need of the parent to conserve resources for any future offspring) is resolved by the reciprocal exchange of information before and after birth. Hormonal signaling ensures a match between parental capacity for resource provisioning and offspring behavior and development after hatching that meet the mother's capacity to provide resources.

Winner and loser effects, in which the outcome of an aggressive encounter influences the tendency to escalate future conflicts, have been documented in many taxa, but we have limited understanding of why they have evolved. One possibility is that individuals use previous victories and defeats to assess their fighting ability relative to others. We explored this idea by modelling a population of strong and weak individuals that do not know their own strength, but keep track of how many fights they have won. Under these conditions, adaptive behaviour generates clear winner and loser effects: individuals who win fights should escalate subsequent conflicts, whereas those who lose should retreat from aggressive opponents. But these effects depend strongly on age and experience. Young, naive individuals should show highly aggressive behaviour and pronounced loser effects. For these inexperienced individuals, fighting is especially profitable because it yields valuable information about their strength. Aggression should then decline as an individual ages and gains experience, with those who lose fights becoming more submissive. Older individuals, who have a better idea of their own strength, should be more strongly influenced by victories than losses. In conclusion, we predict that both aggressiveness and the relative magnitude of winner and loser effects should change with age, owing to changes in how individuals perceive their own strength.