Consolation, i.e., postconflict affiliative interaction directed from a third party to the recipient of aggression, is assumed to have a stress-alleviating function. This function, however, has never been demonstrated. This study shows that consolation in chimpanzees reduces behavioral measures of stress in recipients of aggression. Furthermore, consolation was more likely to occur in the absence of reconciliation, i.e., postconflict affiliative interaction between former opponents. Consolation therefore may act as an alternative to reconciliation when the latter does not occur. In the debate about empathy in great apes, evidence for the stress-alleviating function of consolation in chimpanzees provides support for the argument that consolation could be critical behavior. Consistent with the argument that relationship quality affects their empathic responses, we found that consolation was more likely between individuals with more valuable relationships. Chimpanzees may thus respond to distressed valuable partners by consoling them, thereby reducing their stress levels, especially in the absence of reconciliation.empathy ͉ Pan troglodytes ͉ postconflict behavior ͉ social relationships

A novel approach to studying social relationships in captive adult chimpanzees (Pan troglodytes) was taken by using principal components analysis (PCA) to extract three key components of relationship quality from nine behavioural variables. Based on the loadings of the behavioural variables, the components appeared to match previously hypothesized critical aspects of social relationships and were therefore labelled Value, Compatibility and Security. The effects of kinship, sex combination, age difference and time spent together on each of the relationship quality components were analysed. As expected, kin were found to have more valuable, compatible and secure relationships than non‐kin. Female–female dyads were found to be more compatible than male–male or mixed‐sex dyads, whereas the latter were found to be most secure. Partners of a similar age were found to have more secure and more valuable relationships than those with a larger age gap. Individuals that were together in the group for longer were more valuable and more compatible, but their relationships were found to be less secure than individuals that were together in the group for a shorter time. Although some of the results may be unexpected based on chimpanzee socio‐ecology, they fit well overall with the history and social dynamics of the study group. The methods used confer a significant advantage in producing quantitative composite measures of each component of relationship quality, obtained in an objective manner. These findings therefore promote the use of such measures in future studies requiring an assessment of the qualities of dyadic social relationships.

The quality of a social relationship represents the history of past social interactions between two individuals, from which the nature and outcome of future interactions can be predicted. Current theory predicts that relationship quality comprises three separate components, its value, compatibility and security. This study is the first to investigate the components of relationship quality in a large-brained bird. Following methods recently used to obtain quantitative measures of each relationship quality component in chimpanzees, , we entered data on seven behavioural variables from a group of 11 ravens,, into a principal components analysis. The characteristics of the extracted components matched those predicted for value, compatibility and security, and were labelled as such. When the effects of kinship and sex combination on each relationship quality component were analysed, we found that kin had more valuable relationships, whereas females had less secure and compatible relationships, although the effect of sex combination on compatibility only applied to nonkin. These patterns are consistent with what little knowledge we have of raven relationships from aviary studies and show that the components of relationship quality in ravens may indeed be analogous to those in chimpanzees.

BackgroundBystander affiliation (post-conflict affiliation from an uninvolved bystander to the conflict victim) may represent an expression of empathy in which the bystander consoles the victim to alleviate the victim's distress (“consolation”). However, alternative hypotheses for the function of bystander affiliation also exist. Determining whether ravens spontaneously offer consolation to distressed partners may not only help us to understand how animals deal with the costs of aggressive conflict, but may also play an important role in the empathy debate.Methodology/Principal findingsThis study investigates the post-conflict behavior of ravens, applying the predictive framework for the function of bystander affiliation for the first time in a non-ape species. We found weak evidence for reconciliation (post-conflict affiliation between former opponents), but strong evidence for both bystander affiliation and solicited bystander affiliation (post-conflict affiliation from the victim to a bystander). Bystanders involved in both interactions were likely to share a valuable relationship with the victim. Bystander affiliation offered to the victim was more likely to occur after intense conflicts. Renewed aggression was less likely to occur after the victim solicited affiliation from a bystander.Conclusions/SignificanceOur findings suggest that in ravens, bystanders may console victims with whom they share a valuable relationship, thus alleviating the victims' post-conflict distress. Conversely victims may affiliate with bystanders after a conflict in order to reduce the likelihood of renewed aggression. These results stress the importance of relationship quality in determining the occurrence and function of post-conflict interactions, and show that ravens may be sensitive to the emotions of others.

Reconciliation, a post-conflict affiliative interaction between former opponents, is an important mechanism for reducing the costs of aggressive conflict in primates and some other mammals as it may repair the opponents' relationship and reduce post-conflict distress. Opponents who share a valuable relationship are expected to be more likely to reconcile as for such partners the benefits of relationship repair should outweigh the risk of renewed aggression. In birds, however, post-conflict behavior has thus far been marked by an apparent absence of reconciliation, suggested to result either from differing avian and mammalian strategies or because birds may not share valuable relationships with partners with whom they engage in aggressive conflict. Here, we demonstrate the occurrence of reconciliation in a group of captive subadult ravens (Corvus corax) and show that it is more likely to occur after conflicts between partners who share a valuable relationship. Furthermore, former opponents were less likely to engage in renewed aggression following reconciliation, suggesting that reconciliation repairs damage caused to their relationship by the preceding conflict. Our findings suggest not only that primate-like valuable relationships exist outside the pair bond in birds, but that such partners may employ the same mechanisms in birds as in primates to ensure that the benefits afforded by their relationships are maintained even when conflicts of interest escalate into aggression. These results provide further support for a convergent evolution of social strategies in avian and mammalian species.