How animals respond to conflict provides key insights into the evolution of socio-cognitive and emotional capacities. Evidence from apes has shown that, after social conflicts, bystanders approach victims of aggression to offer stress-alleviating contact behavior, a phenomenon known as consolation. This other-orientated behavior depends on sensitivity to the other's emotional state, whereby the consoler acts to ameliorate the other's situation. We examined post-conflict interactions in bonobos (Pan paniscus) to identify the determinants of consolation and reconciliation. Thirty-six semi-free bonobos of all ages were observed at the Lola ya Bonobo Sanctuary, DR Congo, using standardized Post-conflict/Matched Control methods. Across age and sex classes, bonobos consoled victims and reconciled after conflicts using a suite of affiliative and socio-sexual behaviors including embracing, touching, and mounting. Juveniles were more likely to console than adults, challenging the assumption that comfort-giving rests on advanced cognitive mechanisms that emerge only with age. Mother-reared individuals were more likely to console than orphans, highlighting the role of rearing in emotional development. Consistent with previous studies, bystanders were more likely to console relatives or closely bonded partners. Effects of kinship, affiliation and rearing were similarly indicated in patterns of reconciliation. Nearby bystanders were significantly more likely to contact victims than more distal ones, and consolation was more likely in non-food contexts than during feeding. The results did not provide convincing evidence that bystander contacts served for self-protection or as substitutes for reconciliation. Overall, results indicate that a suite of social, developmental and contextual factors underlie consolation and reconciliation in bonobos and that a sensitivity to the emotions of others and the ability to provide appropriate consolatory behaviors emerges early in development.

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Chimpanzees (Pan troglodytes) and bonobos (P. paniscus) are our closest living relatives, with the human lineage diverging from the Pan lineage only around five to seven Mya, but possibly as early as eight Mya. Chimpanzees and bonobos even share genetic similarities with humans that they do not share with each other. Given their close genetic relationship to humans, both Pan species represent crucial living models for reconstructing our last common ancestor (LCA) and identifying uniquely human features. Comparing the similarities and differences of the two Pan is thus essential for constructing balanced models of human evolution..

Keywords: culture great ape neoteny Pan primate evolution sex difference tool use Chimpanzees, Pan troglodytes, are the most sophisticated tool-users among all nonhuman primates. From an evolutionary perspective, it is therefore puzzling that the tool use behaviour of their closest living primate relative, the bonobo, Pan paniscus, has been described as particularly poor. However, only a small number of bonobo groups have been studied in the wild and only over comparably short periods. Here, we show that captive bonobos and chimpanzees are equally diverse tool-users in most contexts. Our observations illustrate that tool use in bonobos can be highly complex and no different from what has been described for chimpanzees. The only major difference in the chimpanzee and bonobo data was that bonobos of all ageesex classes used tools in a play context, a possible manifestation of their neotenous nature. We also found that female bonobos displayed a larger range of tool use behaviours than males, a pattern previously described for chimpanzees but not for other great apes. Our results are consistent with the hypothesis that the femalebiased tool use evolved prior to the split between bonobos and chimpanzees.