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Edited by Brian Hare and Shinya Yamamoto

This volume includes twelve novel empirical papers focusing on the behaviour and cognition of both captive and wild bonobos ( Pan paniscus). As our species less known closest relative, the bonobo has gone from being little studied to increasingly popular as a species of focus over the past decade. Overall this volume demonstrates how anyone interested in understanding humans or chimpanzees must also know bonobos. Bonobos are not only equal to chimpanzees as our relatives, but they are also unique.

The majority of papers in this volume show that whether you are interested in the evolution of culture and tool use, social relationships and sharing or foraging ecology and cognition, bonobos have a major contribution to make. Four papers provide further evidence that the behaviour and psychology of bonobo females is radically different from that observed in chimpanzees. Foraging behaviour and cognition of bonobos is the focus of three papers that each show important ways that bonobos spatial cognition differs remarkably from chimpanzees. Two papers are relevant to solving the puzzle of why bonobos are expert extractive foragers in captivity but have never been seen using tools to obtain food in the wild.

The articles presented in this volume are previously published in a Special Issue of Behaviour, Volume 152, Parts 3-4 (March 2015).
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T. Furuichi, C. Sanz, K. Koops, T. Sakamaki, H. Ryu, N. Tokuyama and D. Morgan

Abstract

One of the most conspicuous behavioural differences among great apes is the paucity of tool use among wild bonobos (Pan paniscus) in comparison to chimpanzees (Pan troglodytes) who are one of the most prolific and skilled tool users in the animal kingdom. This is in spite of the fact that bonobo tool use repertories are as large and diverse as chimpanzees’ in captive settings. In this study, we compared tool using behaviours and potential drivers of these behaviours in the Wamba bonobo population located in central Democratic Republic of Congo with the Goualougo chimpanzee population of northern Republic of Congo. The tool use repertoire of wild bonobos was comprised of only 13 behaviours, compared to 42 for chimpanzees. However, the number of tool behaviours observed in each study site was similar between bonobos and chimpanzees, and many types of tool use for social, self-grooming/stimulation, and comfort/protection functions were commonly used by both species. A marked difference is that 25 of 42 tool behaviours exhibited by chimpanzees are performed for feeding, in contrast to a single report of bonobos using a leaf sponge to drink water. We examined whether the differences in tool use repertoires can be explained by the necessity, opportunity, relative profitability, or invention hypotheses. We found that habitat composition and fluctuation of fruit production at these two sites were similar, particularly when compared with variation observed between sites within each species. Thus it was unlikely that the necessity hypothesis explains the lack of tool use for feeding in bonobos. Though further study at Wamba is needed, we did not identify any obvious differences in prey availability that would indicate differences in tool using opportunities between the sites. This study could not test the relative profitability hypothesis, and further research is needed on whether tool use is the most efficient means of calorie or protein intake for wild apes. Bonobos at Wamba formed much larger and stable parties than chimpanzees at Goualougo, which was contrary to the prediction by the invention hypothesis. Another explanation is that differences in tool use behaviour between bonobos and chimpanzees might not be explained by the current ecological or social conditions, but rather by circumstances during the Pleistocene Epoch. The observed species differences might also reflect divergent behavioural predispositions, rather than actual differences in cognitive abilities.

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Zanna Clay and Frans B.M. de Waal

Abstract

Sexual contacts are thought to play an important role in regulating social tension in bonobos (Pan paniscus), and are especially common following aggressive conflicts, either between former opponents or involving bystanders. Nevertheless, research on the factors determining post-conflict sexual contacts, their effectiveness in reducing social tension and the nature of post-conflict sexual behaviour is scarce. Here, we collected data on post-conflict affiliative contacts in bonobos occurring between former opponents (reconciliation) and offered by bystanders towards victims (consolation) to investigate the role of sexual contacts in the regulation of aggressive conflicts compared to non-sexual affiliation behaviours. We tested whether post-conflict sexual contacts: (1) alleviate stress, (2) confer reproductive benefits, (3) mediate food-related conflict and (4) repair valuable social bonds. 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. Consolation and reconciliation were both marked by significant increases in the occurrence of sexual behaviours. Reconciliation was almost exclusively characterized by sexual contacts, although consolation was also characterized by increases in non-sexual behaviours, such as embrace. Adults were more likely to engage in post-conflict sexual contacts than younger bonobos. Consistent with the stress-alleviation hypothesis, victims receiving sexual consolatory contact showed significantly lower rates of self-scratching, a marker of stress in primates, compared to receiving non-sexual contact. Post-conflict sexual contacts were not targeted towards valuable social partners and they did not confer obvious reproductive benefits; nor were they used to mediate food-related conflicts. Overall, results highlight the role of sex in regulating tension and social conflicts in bonobos.

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Jeroen M.G. Stevens, Evelien de Groot and Nicky Staes

Abstract

We use Principal Component Analyses (PCA) to describe components of social relationship quality in bonobos. We find a three component structure, with the first two components, labelled Value and Compatibility, closely matching the theoretical constructs as well as components reported for chimpanzees and other primates. The third component differed but was abandoned based on Parallel Analysis. Among bonobos, female–female dyads have higher Value and Compatibility. Relationships between males are characterised by low Value and Compatibility. Dyads that had been housed together for a longer time and maternally related ones also have more valuable relationships, while individuals close in rank have low compatibility. The results confirm the strong bonds among female bonobos, but for the first time can describe how they differ qualitatively from close bonds reported for captive chimpanzee females. We suggest future studies should also include Parallel Analysis to more accurately describe the number of components in relationship quality.

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Heungjin Ryu, David A. Hill and Takeshi Furuichi

Abstract

Perineal sexual skin swelling in relation to menstrual cycle occurs in a variety of primate taxa. However, sexual swelling with exaggerated size and colour is found only in some Old World monkeys and the two Pan species. Although several hypotheses have been proposed (e.g., reliable indicator hypothesis and graded signal hypothesis), it seems unlikely that a single explanation can account for the significance of the sexual swelling in all of these species. Bonobos (Pan paniscus) provide an excellent opportunity for studying sexual swelling since they have the most prolonged maximal swelling periods among primates. In this study we propose a new hypothesis that sexual swelling in female bonobos increases their attractiveness to other females and thereby facilitates affiliative social interaction with them. We found that free-ranging female bonobos with maximal sexual swelling engaged in affiliative social interactions with other females, including genito-genital rubbing, staying in close proximity and grooming, more frequently than females without maximal swelling. These tendencies suggest that females with maximal swelling were attractive to other females. The results also suggest that the benefits of maximal swelling might vary among females depending on their life-history stage. In particular, young females may get more benefits from prolonged maximal swelling through increased grooming reciprocity and staying in close proximity to other females. Thus our study supported the hypothesis that one function of prolonged maximal swelling in bonobos is to increase attractiveness to other females, thereby enhancing affiliative relationships between females in a male-philopatric social system.

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Hare Brian and Yamamoto Shinya

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Jingzhi Tan, Suzy Kwetuenda and Brian Hare

Abstract

Bonobos are the only ape species, other than humans, that have demonstrated prosocial behaviors toward groupmates and strangers. However, bonobos have not been tested in the most frequently used test of prosociality in animals. The current study tested the other-regarding preferences of bonobos in two experiments using the prosocial choice task. In the first experiment subjects preferred a food option that would benefit both themselves and another bonobo. This preference was likely the result of a location bias developed in the pretest since they showed the same preference in the non-social control condition within test sessions. A second experiment was designed to help subjects overcome this bias that might interfere with their social choices. Bonobos again did not prefer to choose the prosocial option. However, results suggest constraints of this paradigm in revealing social preferences. In discussing our results we consider why bonobos show robust prosocial preferences in other paradigms but not here. While others have suggested that such contradictory results might suggest interesting motivational or cognitive differences between humans and non-humans, we propose that the current ‘standard’ paradigm has failed validation due to three methodological constraints. Across the dozens of studies completed few have demonstrated that non-human subjects understand the causal properties of the apparatus, non-social biases quickly develop in inadequately counterbalanced pretests that typically explain subjects’ choices in the test, and even human children found this choice task too cognitively demanding to consistently show prosocial preferences. We suggest it is time to consider switching to a variety of more powerful and valid measures.

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Shinya Yamamoto

Abstract

Food sharing is considered to be a driving force in the evolution of cooperation in human societies. Previously postulated hypotheses for the mechanism and evolution of food sharing, e.g., reciprocity and sharing-under-pressure, were primarily proposed on the basis of meat sharing in chimpanzees. However, food sharing in bonobos has some remarkably different characteristics. Here I report details pertaining to fruit sharing in wild bonobos in Wamba based on 150 events of junglesop fruit sharing between independent individuals. The bonobos, primarily adult females, shared fruit that could be obtained individually without any cooperation or specialized skills. There was no evidence for reciprocal exchange, and their peaceful sharing seems to contradict the sharing-under-pressure explanation. Subordinate females begged for abundant fruit from dominants; this might indicate that they tested the dominants’ tolerance based on social bonds rather than simply begging for the food itself, suggesting existence of courtesy food sharing in bonobos.

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Brian Hare and Shinya Yamamoto

Abstract

This Special Issue of Behaviour includes twelve novel empirical papers focusing on the behaviour and cognition of both captive and wild bonobos (Pan paniscus). As our species less known closest relative, the bonobo has gone from being little studied to increasingly popular as a species of focus over the past decade. We suggest that bonobos are ready to come off the scientific endangered list as a result. This Special Issue is exhibit A in showing that a renaissance in bonobo research is well underway. In this Editorial we review a number of traits in which bonobos and chimpanzees are more similar to humans than they are each other. We show how this means that bonobos provide an extremely powerful test of ideas about human uniqueness as well as being crucial to determining the evolutionary processes by which cognitive traits evolve in apes. This introduction places the twelve empirical contributions within the Special Issue in the larger evolutionary context to which they contribute. Overall this Special Issue demonstrates how anyone interested in understanding humans or chimpanzees must also know bonobos.

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Victoria Wobber and Esther Herrmann

Abstract

Levels of the steroid hormone testosterone have been found to impact diverse features of cognition from spatial memory to decision-making regarding risk, both in humans and other animals. However less is known about whether closely-related species differ in their testosterone-cognition relationships in line with pressures shaping each species’ cognitive evolution. We therefore examined relationships between testosterone and cognition in two-closely related species that differ markedly in their social behaviour, cognition, and patterns of testosterone production: bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). We presented individuals of both species with a battery of 16 cognitive tasks and determined whether performance on these tasks correlated with average testosterone level. We found that among male chimpanzees, high levels of testosterone correlated with higher performance in numerous tasks, including tasks assessing spatial cognition and physical cognitive abilities more broadly. Meanwhile, in male bonobos we found no correlations between testosterone and performance on the cognitive tasks, and found no correlations in females of either species. Building on prior comparative research, these results suggest that bonobos and chimpanzees differ critically in the proximate mechanisms influencing their cognitive capacities, and that in particular the role of testosterone in shaping behaviour and cognition differs dramatically between the two species.