In the analysis of social dominance in groups of animals, linearity has been used by many researchers as the main structural characteristic of a dominance hierarchy. In this paper we propose, alongside linearity, a quantitative measure for another property of a dominance hierarchy, namely its steepness. Steepness of a hierarchy is defined here as the absolute slope of the straight line fitted to the normalized David's scores (calculated on the basis of a dyadic dominance index corrected for chance) plotted against the subjects' ranks. This correction for chance is an improvement of an earlier proposal by de Vries (appendix 2 in de Vries, Animal Behaviour, 1998, 55, 827-843). In addition, we present a randomization procedure for determining the statistical significance of a hierarchy's steepness, which can be used to test the observed steepness against the steepness expected under the null hypothesis of random win chances for all pairs of individuals. Whereas linearity depends on the number of established binary dominance relationships and the degree of transitivity in these relationships, steepness measures the degree to which individuals differ from each other in winning dominance encounters. Linearity and steepness are complementary measures to characterize a dominance hierarchy.

Tolerant food sharing among human foragers can largely be explained by reciprocity. In contrast, food sharing among chimpanzees and bonobos may not always reflect reciprocity, which could be explained by different dominance styles: in egalitarian societies reciprocity is expressed freely, while in more despotic groups dominants may hinder reciprocity. We tested the degree of reciprocity and the influence of dominance on food sharing among chimpanzees and bonobos in two captive groups. First, we found that chimpanzees shared more frequently, more tolerantly, and more actively than bonobos. Second, among chimpanzees, food received was the best predictor of food shared, indicating reciprocal exchange, whereas among bonobos transfers were mostly unidirectional. Third, chimpanzees had a shallower and less linear dominance hierarchy, indicating that they were less despotic than bonobos. This suggests that the tolerant and reciprocal sharing found in chimpanzees, but not bonobos, was made possible by the absence of despotism. To investigate this further, we tested the relationship between despotism and reciprocity in grooming using data from an additional five groups and five different study periods on the main groups. The results showed that i) all chimpanzee groups were less despotic and groomed more reciprocally than bonobo groups, and ii) there was a general negative correlation between despotism and grooming reciprocity across species. This indicates that an egalitarian hierarchy may be more common in chimpanzees, at least in captivity, thus fostering reciprocal exchange. We conclude that a shallow dominance hierarchy was a necessary precondition for the evolution of human-like reciprocal food sharing.

Much of the debate about reciprocity in humans and other primates hinges on proximate mechanisms, or more precisely, the contingency of one service on another. While there is good evidence for long-term statistical contingencies of services given and received in primates, results for short-term behavioral contingencies are mixed. Indeed, as we show here controlled experiments using artificial tasks and explicit turn-taking were unlikely to find short-term effects. We therefore used more naturalistic experiments to test for short-term contingencies of grooming on food sharing and vice versa in one group of chimpanzees and two groups of bonobos. Overall, we found significant effects of grooming on food sharing and vice versa, however, in the chimpanzees these effects disappeared when controlling for longterm characteristics of the dyad including services exchanged over the whole study period. In the bonobos, short-term contingencies remained significant which was likely a consequence of considerable tension surrounding monopolizable food resulting in higher rates of grooming and other affiliative behaviors around sharing sessions. These results are consistent with the fact that previous evidence for shortterm contingency often involved grooming and that long-term contingency is more commonly observed in primates. We propose that long-term contingency is proximately regulated by a 'relationship score' computed through a tally of past interactions which tend to outweigh recent single events. We therefore suggest that future research into the proximate mechanisms of reciprocity should trace the development of such a score by focusing on newly formed dyads with no history of interactions. Much of the debate about reciprocity in humans and other primates hinges on proximate 3 mechanisms, or more precisely, the contingency of one service on another. While there is good 4 evidence for long-term statistical contingencies of services given and received in primates, 5 results for short-term behavioral contingencies are very mixed. Indeed, controlled experiments 6 using artificial tasks and explicit turn-taking were unlikely to find short-term effects, as we show 7here. We therefore used more naturalistic experiments to test for short-term contingencies of 8 grooming on food sharing and vice versa in one group of chimpanzees and two groups of 9 bonobos. Overall, we found significant effects of grooming on food sharing and vice versa, 10 however, in the chimpanzees these effects disappeared when controlling for long-term 11 characteristics of the dyad including services exchanged over the whole study period. In the 12 bonobos, short-term contingencies remained significant which was likely a consequence of 13 considerable tension surrounding monopolizable food resulting in higher rates of grooming and 14 other affiliative behaviors around sharing sessions. These results are consistent with the fact that 15 previous evidence for short-term contingency often involved grooming and that long-term 16 contingency is more c...

To better understand human and chimpanzee personality evolution, we obtained trait ratings of personality for 154 captive bonobos (~80% of the U.S. and European population). We found factors that we labeled Assertiveness, Conscientiousness, Openness, Agreeableness, Attentiveness, and Extraversion. The interrater reliabilities and test-retest reliabilities for these factors were comparable to those found in humans and other species. Using orthogonal targeted Procrustes rotations, we compared the bonobo dimensions with those of three samples of captive chimpanzees. Overall congruence coefficients indicated a fair degree of similarity; at the factor level, there was good evidence for Assertiveness, Conscientiousness, Openness, and Agreeableness in the chimpanzee samples; evidence for Attentiveness and Extraversion was poor. These findings suggest that, as expected given their close phylogenetic relationship, bonobo personality structure resembles chimpanzee personality structure in some respects. However, divergent evolution, perhaps as a result of socioecological differences between bonobos and chimpanzees, also appears to have shaped personality structure in these species.

Bonobos have a reputation as a female-dominated and egalitarian species. We examined the 2 aspects of dominance in 6 captive bonobo groups. Females do not consistently evoke submission from all males in all contexts. Though females occupy the highest-ranking positions in the dominance hierarchy, there are in each group males that obtain rather high ranks and are able to dominate ≥1 female. Thus female dominance is not complete and hierarchies can be better described as nonexclusive female dominance. We studied egalitarianism by measuring linearity and steepness of dominance hierarchies. The hierarchies of all groups are highly linear. Hierarchies among males are steeper than among females. On average, male bonobos are more despotic than females, but females too can have despotic relations, both with other females and with males. Hence one can call bonobos in captivity semidespotic rather than egalitarian.

Social tolerance is a core aspect of primate social relationships with implications for the evolution of cooperation, prosociality and social learning. We measured the social tolerance of bonobos in an experiment recently validated with chimpanzees to allow for a comparative assessment of group-level tolerance, and found that the bonobo group studied here exhibited lower social tolerance on average than chimpanzees in this paradigm. Furthermore, following the Relational Model of de Waal, we investigated whether bonobos responded to an increased potential for social conflict with tolerance, conflict avoidance or conflict escalation, and found that only behaviours indicative of conflict escalation differed across conditions. Taken together, these findings contribute to the current debate over the level of social tolerance of bonobos and lend support to the position that the social tolerance of bonobos may not be notably high compared with other primates.

The human hand is well known for its unique dexterity which is largely facilitated by a highly mobile, long and powerful thumb that enables both tool manufacturing and use, a key component of human evolution. The bonobo (Pan paniscus), the closest extant relative to modern humans together with the chimpanzee (Pan troglodytes), also possesses good manipulative capabilities but with a lower level of dexterity compared with modern humans. Despite the close phylogenetic relationship between bonobos and humans, detailed quantitative data of the bonobo forelimb musculature remains largely lacking. To understand how morphology may influence dexterity, we investigated the functional anatomy of the bonobo hand using a unique sample of eight bonobo cadavers, along with one chimpanzee and one human (Homo sapiens) cadaver. We performed detailed dissections of unembalmed specimens to collect quantitative datasets of the extrinsic and intrinsic hand musculature, in addition to qualitative descriptions of the forelimb muscle configurations, allowing estimation of force-generating capacities for each functional group. Furthermore, we used medical imaging to quantify the articular surface of the trapeziometacarpal joint to estimate the intra-articular pressure. Our results show that the force-generating capacity for most functional groups of the extrinsic and intrinsic hand muscles in bonobos is largely similar to that of humans, with differences in relative importance of the extensors and rotators. The bonobo thumb musculature has a lower force-generating capacity than observed in the human specimen, but the estimated maximal intra-articular pressure is higher in bonobos. Most importantly, bonobos show a higher degree of functional coupling between the muscles of the thumb, index and lateral fingers than observed in humans. It is conceivable that differentiation and individualization of the hand muscles rather than relative muscle development explain the higher level of dexterity of humans compared with that of bonobos.

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