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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.

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.

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.

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.

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.

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.

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?

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.

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.

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’.

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.

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.

Intraspecific cooperation and interspecific mutualism often feature a marked asymmetry in the scope for exploitation. Cooperation may nevertheless persist despite one-sided opportunities for cheating, provided that the partner vulnerable to exploitation has sufficient control over the duration of interaction. The effectiveness of the threat of terminating an encounter, however, depends upon the ease with which both the potential victim and the potential exploiter can find replacement partners. Here, we extend a simple, game-theoretical model of this form of partner control to incorporate variation in the relative abundance of potential victims and exploiters, which leads to variation in the time required for individuals of each type to find a new partner. We show that such market effects have a dramatic influence on the stable level of exploitation (and consequent duration of interaction). As the relative abundance of victims decreases, they become less tolerant to exploitation, terminating encounters earlier (for a given level of exploitation), whereas exploiters behave in a more cooperative manner. As a result, the stable duration of interaction actually increases, despite the decreasing tolerance of the victims. Below a critical level of relative victim abundance, the model suggests that the cost of finding a replacement partner becomes so great that it does not pay to exploit at all.

Recent studies on mammals investigating parent-of-origin-specific effects such as genomic imprinting and maternal effects have demonstrated their impact on short-term measures of fitness, for example offspring growth. However, the long-term fitness consequences of parent-of-origin-specific effects and their role outside the immediate mother–offspring interaction remain largely unexplored. Here, we show that female mice mated to males that inherited the same set of paternal and maternal genes as themselves have a higher reproductive success than females mated to males of reciprocal genotype. Furthermore, we demonstrate that the early social environment experienced by an individual influences its reproductive success. Females raised with unrelated siblings in a mixed litter had a subsequent lower reproductive success than those that were fostered together with all their biological siblings in unmixed litters. Our results highlight the important influence of parent-of-origin-specific effects and conditions in early development on long-term reproductive success in mammals and suggest that parent-of-origin-specific effects may provide the underlying mechanism for beneficial coadaptation between genotypes, for example, in mate choice.

Many mimetic organisms have evolved a close resemblance to their models, making it difficult to discriminate between them on the basis of appearance alone. However, if mimics and models differ slightly in their activity patterns, behavior, or use of microhabitats, the exact circumstances under which a signaler is encountered may provide additional clues to its identity. We employ an optimality model of mimetic discrimination in which signal receivers obtain information about the relative risk of encountering mimics and models by observing an external background cue and flexibly adjust their response thresholds. Although such flexibility on the part of signal receivers has been predicted by theory and is supported by empirical evidence in a range of biological settings, little is known about the effects it has on signalers. We show that the presence of external cues that partly reveal signaler identity may benefit models and harm mimics, harm both, or even benefit both, depending on ecological circumstances. Moreover, if mimetic traits are costly to express, or mimics are related to their neighbors, context-dependent discrimination can dramatically alter the outcome of mimetic evolution. We discuss context-dependent discrimination among signal receivers in relation to small-scale synchrony in model and mimic activity patterns.

The resemblance between mimetic organisms and their models varies from near perfect to very crude. One possible explanation, which has received surprisingly little attention, is that evolution can improve mimicry only at some cost to the mimetic organism. In this article, an evolutionary game theory model of mimicry is presented that incorporates such constraints. The model generates novel and testable predictions. First, Batesian mimics that are very common and/or mimic very weakly defended models should evolve either inaccurate mimicry (by stabilizing selection) or mimetic polymorphism. Second, Batesian mimics that are very common and/or mimic very weakly defended models are more likely to evolve mimetic polymorphism if they encounter predators at high rates and/or are bad at evading predator attacks. The model also examines how cognitive constraints acting on signal receivers may help determine evolutionarily stable levels of mimicry. Surprisingly, improved discrimination abilities among signal receivers may sometimes select for less accurate mimicry.

Most attempts to model the evolution of parental care assume that caring and mating are mutually exclusive activities (i.e., individuals acquire and guard broods "sequentially"). However, in most fish and certain insects, males can keep mating and collecting additional eggs while continuing to guard broods obtained earlier (i.e., males guard "overlapping" broods). We present a model of parental care with overlapping broods in which males can mate and guard simultaneously, even though there is a trade-off between these two activities. Within this framework, we show that male care is favored by short female processing times and high population densities, which minimize the mating cost of care. Relatively low mortality while guarding is also important for the stability of male care. Female care, on the other hand, is favored by long female processing times and low populations densities, which lead to longer intermating intervals. Biparental care is stable only when the cost to benefit ratio of care was not biased toward either sex. We derive quantitative estimates of fitness for different strategies for two species of assassin bugs with male and female uniparental care and show that the model predicts the correct form of care for both species. We believe our model might help explain the prevalence of male uniparental care in certain taxa, such as fish.

Godfray's influential model of competitive begging predicted that offspring should respond to each other's behavior, displaying more intensely when competing with needier rivals. Empirical tests of this prediction have, however, yielded equivocal results. Here, I develop a series of evolutionarily stable strategy models of begging as a signal of need, which show that this prediction holds only for competitive aspects of display that influence the division of food among the brood. No such response is expected for cooperative begging (which influences the total level of provisioning by the parent), and the models even predict the opposite trend under some circumstances (where the indirect costs of extracting additional resources from the parent are high). These contrasting sets of predictions may help to explain the varying empirical results obtained by studies of sibling interaction. Cooperative (as opposed to competitive) begging is likely to be of greater significance in cases where dominant young can gain direct control of allocation (or enjoy some competitive advantage). Dominants are then predicted to parasitize the efforts of their weaker rivals and reduce their own investment in cooperative signaling while continuing to claim a disproportionately large share of the resources provided by the parent.

Theoretical models have demonstrated the possibility of stable cost-free signalling of need between relatives. The stability of these cost-free equilibria depends on the indirect fitness cost of cheating and deceiving a donor into giving away resources. We show that this stability is highly sensitive to the distribution of need among signallers and receivers. In particular, cost-free signalling is likely to prove stable only if there is very large variation in need (such that the least-needy individuals stand to gain much less than the most-needy individuals from additional resources). We discuss whether these conditions are likely to be found in altricial avian breeding systems--the most intensively studied instance of signalling of need between relatives. We suggest that cost-free signalling is more likely to prove stable and will provide parents with more information during the earlier phases of chick growth, when parents can more easily meet the demands of a brood (and chicks are more likely to reach satiation). Later, informative yet cost-free signalling is unlikely to persist.

In an uncertain world, animals face both unexpected opportunities and danger. Such outcomes can select for two potential strategies: collecting information to reduce uncertainty, or insuring against it. We investigate the relative value of information and insurance (energy reserves) under starvation risk by offering model foragers a choice between constant and varying food sources over finite foraging bouts. We show that sampling the variable option (choosing it when it is not expected to be good) should decline both with lower reserves and late in foraging bouts; in order to be able to reap the reduction in uncertainty associated with exploiting a variable resource effectively, foragers must be able to afford and compensate for an initial increase in the risk of an energetic shortfall associated with choosing the option when it is bad. Consequently, expected exploitation of the varying option increases as it becomes less variable, and when the overall risk of energetic shortfall is reduced. In addition, little activity on the variable alternative is expected until reserves are built up early in a foraging bout. This indicates that gathering information is a luxury while insurance is a necessity, at least when foraging on stochastic and variable food under the risk of starvation.

Recent signalling models have shown that honest, cost-free communication between relatives can be stable. Moreover, cost-free signalling equilibria are in some cases more efficient than costly equilibria. However, we show that they are also relatively uninformative, particularly when relatedness between signaller and receiver is low. We explore the trade-off between signal cost and information, and further demonstrate that incorporating competition among signallers into a model of communication between relatives can reduce the propensity of any one signaller to display. As a result, there is a general increase in the amount of broadcast information in a non-costly signal with increasing competitor number.

One key trade-off underlying life-history evolution is the one between age and size at maturity, with earlier maturation leading to greater chances of juvenile survival at the cost of reduced fecundity as an adult. Here we model the impact of limited dispersal and kin competition on the stable resolution of this trade-off. We show that if mating is at least occasionally nonlocal, then limited dispersal favors juvenile survival over adult fecundity in females, promoting earlier female maturation at the population level; at the same time, it favors adult fecundity over juvenile survival in males, promoting later male maturation. Limited dispersal and local competition can thus drive the evolution of sexual dimorphism in the timing of maturation and consequent dimorphism in body size. At the individual level, if maturation can be flexibly adjusted in response to dispersal status, then both males and females who disperse as offspring should mature earlier than those who remain on their natal patch.

The evolution of cooperation in animal and human societies is associated with mechanisms to suppress individual selfishness. In insect societies, queens and workers enforce cooperation by “policing” selfish reproduction by workers. Insect policing typically takes the form of damage limitation after individuals have carried out selfish acts (such as laying eggs). In contrast, human policing is based on the use of threats that deter individuals from acting selfishly in the first place, minimizing the need for damage limitation. Policing by threat could in principle be used to enforce reproductive suppression in animal societies, but testing this idea requires an experimental approach to simulate reproductive transgression and provoke out-of-equilibrium behavior. We carried out an experiment of this kind on a wild population of cooperatively breeding banded mongooses (Mungos mungo) in Uganda. In this species, each group contains multiple female breeders that give birth to a communal litter, usually on the same day. In a 7-y experiment we used contraceptive injections to manipulate the distribution of maternity within groups, triggering hidden threats of infanticide. Our data suggest that older, socially dominant females use the threat of infanticide to deter selfish reproduction by younger females, but that females can escape the threat of infanticide by synchronizing birth to the same day as older females. Our study shows that reproduction in animal societies can be profoundly influenced by threats that remain hidden until they are triggered experimentally. Coercion may thus extend well beyond the systems in which acts of infanticide are common.

Some individuals (helpers) in cooperatively breeding species provide alloparental care and often suppress their own reproduction. Kin selection is clearly an important explanation for such behaviour, but a possible alternative is group augmentation where individuals survive or reproduce better in large groups and where it therefore pays to recruit new members to the group. The evolutionary stability of group augmentation is currently disputed. We model evolutionarily stable helping strategies by following the dynamics of social groups with varying degrees of subordinate help. We also distinguish between passive augmentation, where a group member benefits from the mere presence of others, and active augmentation, where their presence as such is neutral or harmful, but where helping to recruit new group members may still be beneficial if they in turn actively provide help for the current reproductives ('delayed reciprocity'). The results show that group augmentation (either passive or active) can be evolutionarily stable and explain costly helping by non-reproductive subordinates, either alone or leading to elevated help levels when acting in concert with kin selection. Group augmentation can thus potentially explain the weak relationships between relatedness and helping behaviour that are observed in some cooperatively breeding species. In some cases, the superior mutualistic performance of cooperatively behaving groups can generate an incentive to stay and help which is strong enough to make ecological constraints unnecessary for explaining the stability of cooperatively breeding groups.

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 many instances of cooperation, only one individual has both the potential and the incentive to ‘cheat’ and exploit its partner. Under these asymmetric conditions, a simple model predicts that variation in the temptation to cheat and in the potential victim's capacity for partner control leads to shifts between exploitation and cooperation. Here, we show that the threat of early termination of an interaction was sufficient to induce cleaner wrasse Labroides dimidiatus to feed selectively against their preference (which corresponds to cooperatively eating client fish ectoparasites), provided that their preference for alternative food was weak. Under opposite conditions, cleaners fed selectively according to their own preference (which corresponds to cheating by eating client mucus). By contrast, a non-cleaning fish species, Halichoeres melanurus , failed to adjust its foraging behaviour under these same conditions. Thus, cleaners appear to have evolved the power to strategically adjust their levels of cooperation according to the circumstances.

Conflicts of interest arise between signaller and receiver in most kinds of biological communication. Some authors have argued that this conflict is likely to give rise to deceit and exploitation, as receivers lag behind in the coevolutionary ‘arms race’ with signallers. Others have argued that such manipulation is likely to be short-lived and that receivers can avoid being deceived by paying attention to signals that are costly and hence ‘unfakeable’. These two views have been hard to reconcile. Here, we present results from simulations of signal evolution using artificial neural networks, which demonstrate that honesty can coexist with a degree of exploitation. Signal cost ensures that receivers are able to obtain some honest information, but is unable to prevent exploitative signalling strategies from gaining short-term benefits. Although any one receiver bias that is open to exploitation will subsist for only a short period of time once signallers begin to take advantage of it, new preferences of this kind are constantly regenerated through selection and random drift. Hidden preferences and sensory exploitation are thus likely to have an enduring influence on the evolution of honest, costly signals. At the same time, honesty and cost are prerequisites for the evolution of exploitation. When signalling is cost-free, selection cannot act to maintain honesty, and receivers rapidly evolve to ignore signals. This leads to a reduction in the extent of hidden preference, and a consequent loss of potential for exploitation.

SUMMARY (i) To find out whether a mating preference could have initially evolved for adaptive reasons, one must determine whether the preferred trait could have provided useful information about mate quality at the time when the preference first arose. (ii) One way to do so is to determine whether the preference evolved before or after the preferred trait. If the preference evolved first, then it cannot initially have served an adaptive function in mate choice, rather it must have arisen by random drift, or as a pleiotropic consequence of selection acting on other aspects of individual perceptual abilities. (iii) A number of studies have shown that females exhibit a mating preference (e.g. for movement) in non-sexual contexts also, which suggests that it may have evolved for reasons unconnected to mate choice. In addition, phylogenetic analyses have revealed that in several cases, females of a certain taxon exhibit a preference for a male trait that is absent in a sister taxon and in outgroup taxa, and that this preference is shared by females of the sister taxon tacking the male trait. The principle of parsimony suggests that such a preference has been inherited from a common ancestor, while the preferred trait arose only once in the lineage exhibiting the trait, i.e. that the preference predates the attractive trait. (iii) While the above evidence indicates that females may possess ‘hidden’ preferences for male traits that are not exhibited by members of their own species, and that in at least some cases males have later evolved display traits that exploit preexisting preferences of this kind, there have been too few historical studies of preference evolution to allow one to assess the frequency of such exploitation. Moreover historical studies cannot provide strong support for the adaptive origin hypothesis, because coevolution of trait and preference (as opposed to exploitation of a pre-existing bias) is compatible with Fisherian models of preference evolution as well as with honest advertisement and the handicap principle. One can conclude only that while some mating preferences did not originally evolve for adaptive reasons, others may or may not have done so. (iv) To find out whether a mating preference is currently maintained by natural selection because the preferred trait provides useful information about mate quality, one must investigate the phenotypic and genotypic correlates of display, and the fitness consequences of mate choice. (v) A review of the published data reveals some support for the ideas of adaptive choice and honest advertisement. In a number of species, preferred display traits are correlated with putative measures of quality, and in a small proportion of these, there is evidence that reproductive success and/or offspring performance are higher for individuals mated to attractive partners. Very few studies report a failure to find any such correlates of display or any such benefits. (vi) While the above result suggests that honest advertisement does sometimes occur in extant populations (which does not necessarily imply that preferred traits originally evolved as reliable indicators of mate quality), the possibility of publication bias means that one cannot assess how widespread it is. More data is needed to remedy this problem, particularly regarding the fitness consequences of mate choice for females. Experimental rather than observational methods are the best means to gather such data. Studies that look for correlates of display, for instance, should rely on experimentally induced rather than natural variation in ‘quality’. (vii) The most common correlates of male display are age and dominance. The latter observation suggests that there may often be interactions between the processes of intersexual and intrasexual selection. (viii) There is considerably more evidence to support the idea of female choice for direct than for indirect benefits. At the same time, however, it is apparent that mating decisions are commonly influenced by more than one measure of quality, so that these two kinds of choice need not be independent. To assess this possibility will require more studies of the relationship between male attractiveness and offspring performance. (ix) Mate choice is frequently based on more than one display trait, and each trait is frequently influenced by more than one aspect of quality. ‘One quality, one trait’ views of honest advertisement are simplistic, and must be abandoned. (x) Honesty in sexual displays is sometimes maintained by cost (as in strategic handicap models) and sometimes, with approximately equal frequency, by physical necessity (as in revealing handicap models). In some cases, both mechanisms are involved in a single signalling system. To further distinguish between these possibilities will require experimental investigation of display cost, based on manipulation of display traits.

Asymmetries in the costs and benefits of parental investment for mothers, fathers and offspring result in family conflict over the production and provisioning of young1,2,3. In species where females provide most resources before and after birth, the resolution of this conflict may be influenced by genes expressed in mothers and by maternally and paternally inherited genes expressed in offspring4,5. Here we disentangle these effects by means of reciprocal mating and cross-fostering of litters between two strains of mice that differ with respect to the typical resolution of family conflict. We find that differences in litter size between these two strains are determined by paternal genotype, whereas differences in provisioning are under maternal control, showing that there is antagonistic coadaptation of maternal and paternal effects on distinct life-history traits. Maternal provisioning is also influenced by the type of foster offspring. Contradictory to theoretical expectations, however, we find no evidence for a negative correlation across strains between maternal provisioning and offspring demand. Instead, we show that there is positive coadaptation such that offspring obtain more resources from foster mothers of the same strain as their natural mother, irrespective of their father's strain.

Identifying the factors that modulate cooperative and competitive behaviours is the key to understanding social evolution. However, how ecological factors affect social conflict and their fitness consequences remain relatively unexplored. Here, using both a game-theoretical model and empirical data, we show that Taiwan yuhinas (Yuhina brunneiceps)--a joint-nesting species in which group members are unrelated--employ more cooperative strategies in unfavourable environmental conditions. Fighting duration was lower, fewer total eggs were laid and incubation was more likely to start after all females completed egg laying (which causes more synchronous egg hatching). Surprisingly, as a consequence, there were more surviving offspring in unfavourable conditions because the cooperative strategies resulted in fewer dead nestlings. To our knowledge, this study is the first theoretical analysis and empirical study demonstrating that an unfavourable environment reduces social conflict and results in better fitness consequences in social vertebrates.

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.

Studies of coordinated movement have found that, in many animal species, bolder individuals are more likely to initiate movement and shyer individuals to follow. Here, we show that in pairs of foraging stickleback fish, leadership is not merely a passive consequence of temperamental differences. Instead, the act of initiating a joint foraging trip out of cover itself brings about a change in the role that an individual plays throughout the subsequent trip, and success in recruiting a partner affects an individual's tendency to initiate the next trip. On each joint trip, whichever fish took the initiative in leading out of cover gains greater influence over its partner's behaviour, which persists even after several changes in position (i.e. termination attempts and re-joining). During any given trip, the initiator is less responsive to its partner's movements than during trips initiated by the partner. An individual's personality had an important effect on its response to failure to recruit a partner: while bold fish were unaffected by failures to initiate a joint trip, shy individuals were less likely to attempt another initiation after a failure. This difference provides a positive feedback mechanism that can partially stabilise social roles within the pair, but it is not strong enough to prevent occasional swaps, with individuals dynamically adjusting their responses to one another as they exchange roles.

The evolution of group living has attracted considerable attention from behavioral ecologists working on a wide range of study species. However, theoretical research in this field has been largely focused on cooperative breeders. We extend this line of work to species that lack alloparental care (hereafter termed "noncooperative species") but that may benefit from grouping by jointly defending a common territory. We adopt a demographically explicit approach in which the rates of births and deaths as well as the dispersal decisions of individuals in the population determine the turnover rates of territories and the competition for breeding vacancies thus arising. Our results reveal that some of the factors thought to affect the evolution of cooperative breeding also affect the evolution of group living in noncooperative species. Specifically, high fecundity and low mortality of resident individuals both increase the degree of habitat saturation and make joining an established group more profitable for nonresidents (floaters). Moreover, if floaters can forcefully take over territories, the degree of habitat saturation also affects the chance that residents become targets of takeovers. In this situation, communal defense of territories becomes an important benefit that further promotes the evolution of group living.

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.

The evolution of group-living has fascinated but also puzzled researchers from the inception of behavioural ecology. We use a simple optimality approach to examine some of the costs and benefits of group-living in redfronted lemurs ( Eulemur fulvus rufus ). We show that dominant males profit from accepting subordinates within their groups, as the latter significantly decrease the likelihood that the group is taken over by intruders. This benefit is large enough to outweigh the costs of reproductive competition and may constitute the driving force behind the evolution of multi-male associations in this species.

The amount of DNA in the nuclear genome (the DNA C-value) of eukaryotes varies at least 80,000-fold across species, and yet bears little or no relation to organismic complexity or to the number of protein-coding genes. This phenomenon is known as the C-value paradox. One explanation for the C-value paradox attributes the size of the nuclear genome to 'junk' (typically non-coding) genetic elements that accumulate until the costs to the organism of replicating excess DNA select against it. Across species, organisms that develop at a slower rate should tolerate more junk DNA. Alternatively, junk DNA may function as a nucleo-skeleton to maintain the volume of the nucleus at a size proportional to the volume of the cytoplasm in the cell. Across species, the DNA C-value is predicted to vary with the nuclear and cytoplasmic volumes of cells. Previous studies have not been able to distinguish between the skeletal-DNA and junk-DNA explanations for the C-value paradox. We report a study of DNA content in 24 salamander species which does. The size of the nuclear genome is correlated with developmental rate even after the effects of nuclear and cytoplasmic volume have been removed. However, genome size is not correlated with cytoplasmic volume after controlling for developmental rate. These results support the view that junk DNA accumulates in the nuclear genome until the costs of replicating it become too great, rather than that it functions as a nucleo-skeleton.

An analysis of Maynard Smith's two-player, ESS model of biological signalling, the "Sir Philip Sidney game", is presented. The stable strategies of the players in this game are shown to satisfy the conditions of Zahavi's handicap principle. At equilibrium, signals are honest, costly, and costly in a way that is related to the true quality revealed. Further analysis reveals that the level of cost required to maintain stability is inversely related to the degree of relatedness between the players. It therefore seems likely that stable biological signalling systems will feature lower signalling costs when communication occurs between relatives. A three-player, extended version of the model is investigated, in which signals are passed via an intermediate, or "messenger". It is shown that this destabilizes the signalling system, and leads to increased signalling costs. This result suggests that "kin conflict" theories of the evolution of the endosperm in flowering plants require further refinement. The introduction of a novel resource acquisition tissue, which mediates parent-offspring interaction during development, cannot be assumed to limit parent-offspring conflict simply because it carries an extra copy of the maternally inherited genes. The ability to add such complications to the Sir Philip Sidney game and still obtain solutions makes it a very useful modelling tool.

The handicap theory of sexual selection suggests that females prefer mates who display extravagant ornaments that advertise their quality or condition. It is often assumed that as such ornamental traits undergo sexually-selected exaggeration, they must inevitably become more sensitive to condition, and thus more informative. Here, we show that this is not necessarily the case. Depending on the precise form of the relationship between trait size and cost, expression may become more or less condition-dependent as the trait undergoes exaggeration, or may remain unchanged. This leads us to question how much of the information content of sexual signals can be attributed to sexual selection, and how much to pre-existing, naturally-selected condition-dependence.

SummaryIn many animal groups, certain individuals consistently appear at the forefront of coordinated movements [1–4]. How such leaders emerge is poorly understood [5, 6]. Here, we show that in pairs of sticklebacks, Gasterosteus aculeatus, leadership arises from individual differences in the way that fish respond to their partner's movements. Having first established that individuals differed in their propensity to leave cover in order to look for food, we randomly paired fish of varying boldness, and we used a Markov Chain model to infer the individual rules underlying their joint behavior. Both fish in a pair responded to each other's movements—each was more likely to leave cover if the other was already out and to return if the other had already returned. However, we found that bolder individuals displayed greater initiative and were less responsive to their partners, whereas shyer individuals displayed less initiative but followed their partners more faithfully; they also, as followers, elicited greater leadership tendencies in their bold partners. We conclude that leadership in this case is reinforced by positive social feedback.

In a spatially structured population, limited dispersal gives rise to local relatedness, potentially favoring indiscriminate helping behavior. However, it also leads to local competition, which reduces the benefits of helping local kin. This tension has become the focus for a growing body of theoretical work. Existing models, however, have focused chiefly on the net impact of limited dispersal on cooperative or competitive effort in a homogeneous population. Here, I extend existing models of kin selection in a group-structured population to allow for asymmetries in expected fecundity and reproductive success among group members. I explore the consequent impact of limited dispersal on the evolution of helping and harming behavior, and on the degree of reproductive inequality or skew. I show that when individuals in a group differ in their expected fecundity, limited dispersal gives rise to kin selection for harming behavior on the part of more fecund individuals, and for helping behavior on the part of less fecund individuals. As a result, philopatry tends to exaggerate differences in reproductive success, and so promotes greater reproductive skew.

In this article, we explore the impact of sex-biased dispersal on local relatedness and on selection for helping and harming behavior among males and females. We show that in a patch-structured population, when there is a marked sex bias in dispersal, selection will almost always favor harming behavior among individuals of the sex more prone to dispersal. This result holds regardless of the effects of mating skew or overlapping generations. Selection may well also favor helping behavior among individuals of the philopatric sex, particularly if there is generational overlap, but this is less likely to occur if individuals of the philopatric sex compete more intensely for fewer breeding opportunities. In this last case, if generational overlap is low and mating skew pronounced, the result may be selection for harming behavior among both males and females. In general, the rate of dispersal and the level of relatedness among individuals of one sex do not reliably predict their level of helping or harming behavior; selection on either males or females depends on the dispersal of both sexes.

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.

Intraspecific cooperation and interspecific mutualism often feature an asymmetry in the scope for exploitation. We investigate the evolution of indirect reciprocity in an asymmetric game, loosely modelled on interactions between cleaner fishes and clients, in which ‘actors’ can choose to help or to exploit a ‘recipient’ that approaches them, while recipients can only choose whether or not to approach an actor (based on the observation of its behaviour towards others). We show that when actors vary in state over time, in a manner that influences the potential gains from exploitation, an equilibrium is possible at which recipients avoid actors whom they have observed exploiting others in the past, and actors help when the potential gains from exploitation are low but choose to exploit when the potential gains are high. In this context, helping is favoured not because it elicits reciprocal altruism (‘help so that you may be helped’), but because it facilitates profitable exploitation (‘help so that you may gain the opportunity to harm’). The cost of helping one recipient is thereby recouped through exploitation of another. Indirect reciprocity is thus possible even in asymmetric interactions in which one party cannot directly ‘punish’ exploitation or ‘reward’ helping by the other.

We present a synopsis about the commentaries to the target article “Integrating cooperative breeding into theoretical concepts of cooperation”, in which we attempted to integrate general mechanisms to explain cooperative behaviour among unrelated individuals with classic concepts to explain helping behaviour in cooperative breeders that do not invoke kin-based benefits. Here we (1) summarize the positions of the commentators concerning the main issues we raised in the target article and discuss important criticisms and extensions. (2) We relate our target article to some recent reviews on the evolution of cooperation and, (3) clarify how we use terminology with regard to cooperation and cooperative behaviour. (4) We discuss several aspects that were raised with respect to cooperative interactions including by-product mutualism, generalised reciprocity and multi-level selection and, (5) examine the alternatives to our classification scheme as proposed by some commentaries. (6) Finally, we highlight several aspects that might hinder the application of game theoretical mechanisms of cooperation in cooperatively breeding systems. Although there is broad agreement that cooperative breeding theory should be integrated within the more general concepts of cooperation, there is some debate about how this may be achieved. We conclude that the contributions in this special issue provide a fruitful first step and ample suggestions for future directions with regard to a more unified framework of cooperation in cooperative breeders.