'Social competence' refers to the ability of an individual to optimise its social behaviour depending on available social information. Although such ability will enhance social interactions and thus raise Darwinian fitness, its evolutionary and ecological significance has been largely ignored. Social competence is based on behavioural flexibility. We propose that the study of social competence requires an integrative approach that aims to understand how the brain translates social information into flexible behavioural responses, how flexibility might be constrained by the developmental history of an individual or by trade-offs with other (ecological) competences, and how social plasticity feeds back on fitness. Finally we propose a hypothesis of how social competence can become a driver of social evolution.

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There is increasing evidence that the environment experienced early in life can strongly influence adult life histories. It is largely unknown, however, how past and present conditions influence suites of life-history traits regarding major life-history trade-offs. Especially in animals with indeterminate growth, we may expect that environmental conditions of juveniles and adults independently or interactively influence the life-history trade-off between growth and reproduction after maturation. Juvenile growth conditions may initiate a feedback loop determining adult allocation patterns, triggered by size-dependent mortality risk. I tested this possibility in a long-term growth experiment with mouthbrooding cichlids. Females were raised either on a high-food or low-food diet. After maturation half of them were switched to the opposite treatment, while the other half remained unchanged. Adult growth was determined by current resource availability, but key reproductive traits like reproductive rate and offspring size were only influenced by juvenile growth conditions, irrespective of the ration received as adults. Moreover, the allocation of resources to growth versus reproduction and to offspring number versus size were shaped by juvenile rather than adult ecology. These results indicate that early individual history must be considered when analysing causes of life-history variation in natural populations.

When organisms encounter environments that are heterogeneous in time, phenotypic plasticity is often favored by selection. The degree of such plasticity can vary during an organism's lifetime, but the factors promoting differential plastic responses at different ages or life stages remain poorly understood. Here we develop and analyze an evolutionary model to investigate how environmental information is optimally collected and translated into phenotypic adjustments at different ages. We demonstrate that plasticity must often be expected to vary with age in a non-monotonic fashion. Early in life it is generally optimal to delay phenotypic adjustments until sufficient information has been collected about the state of the environment to warrant a costly phenotypic adjustment. Towards the end of life, phenotypic adjustments are disfavored as well, because their beneficial effects can no longer fully be reaped before death.Our analysis clarifies how patterns of age-dependent plasticity are shaped by the interplay of environmental uncertainty, the accuracy of perceived information and the costs of phenotypic adjustments with life-history determinants such as the relative strengths of fecundity and viability selection experienced by the organism over its lifetime. We conclude by comparing our results with expectations for alternative mechanisms, including developmental constraints, that promote age-dependent plasticity.3

Through non-genetic maternal effects, mothers can tailor offspring phenotype to the environment in which young will grow up. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring's environment than the adult environment of mothers. In this case maternal decisions about investment in offspring quality should already be determined during the juvenile phase of mothers. I tested this hypothesis by manipulating juvenile and adult maternal environments independently in a cichlid fish. Females raised in a poor environment produced larger young than females raised without food limitations, irrespective of the feeding conditions experienced during adulthood. This maternal boost was due to a higher investment in eggs and to faster larval growth. Apparently, mothers prepare their offspring for similar environmental conditions to those they encountered as juveniles. This explanation is supported by the distribution of these fishes under natural conditions. Juveniles live in a different and much narrower range of habitats than adults. Therefore, the habitat mothers experienced as juveniles will allow them to predict their offspring's environment better than the conditions in the adult home range.

Social competence is defined as the ability of an animal to optimize the expression of social behaviour as a function of the available social information. The social environment encountered early in life can affect the expression of various social behaviours later in life. We investigated whether early social experience can affect social competence. In the cooperatively breeding cichlid Neolamprologus pulcher, we tested whether individuals reared with older brood-caring conspecifics persistently perform better in a series of tasks (1) simulating different social contexts, (2) assigning individuals different social roles and (3) exposing them to an unknown social situation. Fish that had been reared together with older conspecifics showed more appropriate behaviours both as winners (more aggressive displays) and as losers (more submissive displays) when aggressively competing with peers over a resource, and when trying to be accepted as subordinate group member and prospective brood care helper by an unfamiliar dominant pair (more submissive displays near shelters), a situation they had never encountered before. In both tasks fish that had grown up with older fish were tolerated better by conspecifics than fish reared with same-age siblings only. We detected effects of the early environment on social behaviour in the juvenile and adult stages of the test fish. Our results suggest that growing up in more complex social groups fosters a general social ability (i.e. social competence) in N. pulcher that improves their performance across different social roles and contexts, and which may provide fitness benefits.

In cooperative breeding systems, dominant breeders sometimes tolerate unrelated individuals even if they inflict costs on the dominants. According to the 'pay-to-stay' hypothesis, (i) subordinates can outweigh these costs by providing help and (ii) dominants should be able to enforce help by punishing subordinates that provide insufficient help. This requires that dominants can monitor helping and can recognize group members individually. In a field experiment, we tested whether cooperatively breeding cichlid Neolamprologus pulcher subordinates increase their help after a forced 'idle' period, how other group members respond to a previously idle helper, and how helper behaviour and group responses depend on group size. Previously idle helpers increased their submissiveness and received more aggression than control helpers, suggesting that punishment occurred to enforce help. Subordinates in small groups increased their help more than those in large groups, despite receiving less aggression. When subordinates were temporarily removed, dominants in small groups were more likely to evict returning subordinates. Our results suggest that only in small groups do helpers face a latent threat of punishment by breeders as predicted by the pay-to-stay hypothesis. In large groups, cognitive constraints may prevent breeders from tracking the behaviour of a large number of helpers.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. abstract: Social animals can greatly benefit from well-developed social skills. Because the frequency and diversity of social interactions often increase with the size of social groups, the benefits of advanced social skills can be expected to increase with group size. Variation in social skills often arises during ontogeny, depending on early social experience. Whether variation of social-group sizes affects development of social skills and related changes in brain structures remains unexplored. We investigated whether, in a cooperatively breeding cichlid, early group size (1) shapes social behavior and social skills and (2) induces lasting plastic changes in gross brain structures and (3) whether the development of social skills is confined to a sensitive ontogenetic period. Rearing-group size and the time juveniles spent in these groups interactively influenced the development of social skills and the relative sizes of four main brain regions. We did not detect a sensitive developmental period for the shaping of social behavior within the 2-month experience phase. Instead, our results suggest continuous plastic behavioral changes over time. We discuss how developmental effects on social behavior and brain architecture may adaptively tune phenotypes to their current or future environments.