Groups Of Captive Chimpanzees Research Articles

Naïve, adult, captive chimpanzees do not socially learn how to make and use sharp stone tools

Although once regarded as a unique human feature, tool-use is widespread in the animal kingdom. Some of the most proficient tool-users are our closest living relatives, chimpanzees. These repertoires however consist primarily of tool use, rather than tool manufacture (for later use). Furthermore, most populations of chimpanzees use organic materials, such as sticks and leaves, rather than stones as tools. This distinction may be partly ecological, but it is also important as chimpanzees are often used as models for the evolution of human material culture, the oldest traces of which consist of manufactured sharp stone tools (so-called “flakes”). Thus, examining the conditions (if any) under which chimpanzees may develop flake manufacture and use can provide insight into the drivers of these behaviours in our own lineage. Previous studies on non-human apes’ ability to make and use flakes focused on enculturated apes, giving them full demonstrations of the behaviour immediately, without providing social information on the task in a stepwise manner. Here we tested naïve, captive chimpanzees (N = 4; three potentially enculturated and one unenculturated subject) in a social learning experimental paradigm to investigate whether enculturated and/or unenculturated chimpanzees would develop flake making and use after social information of various degrees (including a human demonstration) was provided in a scaffolded manner. Even though social learning opportunities were provided, neither the unenculturated subject nor any of the potentially enculturated subjects made or used flakes, in stark contrast to previous studies with enculturated apes. These data suggest that flake manufacture and use is outside of our tested group of captive chimpanzees’ individual and social learning repertoires. It also suggests that high levels of enculturation alongside human demonstrations (and/or training) may be required before captive chimpanzees can develop this behaviour.

Bone-related behaviours of captive chimpanzees (Pan troglodytes) during two excavating experiments

After stone tools, bone tools are the most abundant artefact type in the Early Pleistocene archaeological record. That said, they are still relatively scarce, which limits our understanding of the behaviours that led to their production and use. Observations of extant primates constitute a unique source of behavioural data with which to construct hypotheses about the technological forms and repertoires exhibited by our hominin ancestors. We conducted two different experiments to investigate the behavioural responses of two groups of captive chimpanzees (Pan troglodytes; n = 33 and n = 9) to disarticulated, defleshed, ungulate bones while participating in a foraging task aimed at eliciting excavating behaviour. Each chimpanzee group was provided with bone specimens with different characteristics, and the two groups differed in their respective experience levels with excavating plant tools. We found that several individuals from the inexperienced group used the provided bones as tools during the task. In contrast, none of the individuals from the experienced group used bones as excavating tools, but instead continued using plant tools. These chimpanzees also performed non-excavating bone behaviours such as percussion and tool-assisted extraction of organic material from the medullary cavity. Our findings serve as a proof-of-concept that chimpanzees can be used to investigate spontaneous bone tool behaviours such as bone-assisted excavation. Furthermore, our results raise interesting questions regarding the role that bone characteristics, as well as previous tool-assisted excavating experience with other raw materials, might have in the expression of bone tool-assisted excavation.

Temporal calling patterns of a captive group of chimpanzees (Pan troglodytes)

Temporal calling patterns of a captive group of chimpanzees (Pan troglodytes)

A Comparison of Focal and Opportunistic Sampling Methods when Studying Chimpanzee Facial and Gestural Communication

Researchers frequently use focal individual sampling to study primate communication. Recent studies of primate gestures have shown that opportunistic sampling offers benefits not found in focal individual sampling, such as the collection of larger sample sizes. What is not known is whether the opportunistic method is biased towards certain signal types or signalers. Our goal was to assess the validity of the opportunistic method by comparing focal individual sampling to opportunistic sampling of facial and gestural communication in a group of captive chimpanzees (Pan troglodytes). We compared: (1) the number of observed facial and gestural signals per signal type and (2) the number of observed facial and gestural signals produced by each signaler. Both methods identified facial signals, gesture signals, and gesture signalers at similar relative rates, but the opportunistic sampling method yielded a more even distribution of signalers and signal types than the focal individual sampling method. In addition, the opportunistic sampling method resulted in larger sample sizes for both facial and gestural communication. However, the opportunistic method did not allow us to calculate the signals per time for each individual, which is easily done with the focal individual method. These results suggest that the opportunistic sampling method is (1) comparable to the focal individual sampling method in multiple important measures, (2) associated with additional sampling benefits, and (3) limited in measuring some variables. Thus, we recommend that future studies use a mixed-methods approach, as focal individual and opportunistic sampling have distinct strengths that complement each other’s limitations.

Does group size matter? Captive chimpanzee (Pan troglodytes) behavior as a function of group size and composition.

The National Institutes of Health and the Association of Zoos and Aquariums recommend that captive chimpanzees be housed in multi-male, multi-female, age-diverse groups of no less than seven individuals. These recommendations are rooted in the idea that captive chimpanzee groups should be modeled after free-ranging, wild, fission-fusion chimpanzee societies. However, captive chimpanzees do not face the environmental pressures faced by wild chimpanzees, including food scarcity, inter-group competition, and predation. As such, it has been posited that wild, natural conditions may not be the most relevant metric for defining optimal captive chimpanzee group sizes and compositions. Additionally, captive housing poses a set of restrictions on group sizes and compositions, including the need to balance large, multi-male groups with space per animal limitations and intra-group aggression. In the present study, we examined the behavioral effects of group size, within-group age range, and percentage of males in the group. We collected 713 hr of focal animal samples across 120 captive chimpanzees housed in social groups of 4-10 individuals using a 58-behavior ethogram. Chimpanzees housed in groups with a large age range exhibited less inactivity and more locomotion than chimpanzees housed in groups with smaller age ranges. Additionally, chimpanzees in groups of ≥7 with less than half males showed the highest levels of locomotion. Lastly, chimpanzees in groups of ≥7 with at least half males showed the highest levels of affiliation. There were no other significant differences in behavior as a function of these variables or their interactions. These findings lend some support to the existing group size and composition recommendations, providing empirical evidence that there may be certain advantages to housing captive chimpanzees in larger groups with a more diverse age range and/or more males. These results also have practical implications for behavioral management programs across captive settings.

Comparative Investigations of Social Context-Dependent Dominance in Captive Chimpanzees (Pan troglodytes) and Wild Tibetan Macaques (Macaca thibetana)

Theoretical definitions of dominance, how dominance is structured and organized in nature, and how dominance is measured have varied as investigators seek to classify and organize social systems in gregarious species. Given the variability in behavioral measures and statistical methods used to derive dominance rankings, we conducted a comparative analysis of dominance using existing statistical techniques to analyze dominance ranks, social context-dependent dominance structures, the reliability of statistical analyses, and rank predictability of dominance structures on other social behaviors. We investigated these topics using behavioral data from captive chimpanzees (Pan troglodytes) and wild Tibetan macaques (Macaca thibetana). We used a combination of all-occurrence, focal-animal, and instantaneous scan sampling to collect social, agonistic, and associative data from both species. We analyzed our data to derive dominance ranks, test rank reliability, and assess cross-context predictability using various statistical analyses. Our results indicate context-dependent dominance and individual social roles in the captive chimpanzee group, one broadly defined dominance structure in the Tibetan macaque group, and high within-context analysis reliability but little cross-context predictability. Overall, we suggest this approach is preferable over investigations of dominance where only a few behavioral metrics and statistical analyses are utilized with little consideration of rank reliability or cross-context predictability.

The presence of females induces elevated cortisol levels in an alpha male: Experimental evidence in chimpanzees.

In group-living primates, it has been reported that the alpha male exhibits high concentrations of cortisol and testosterone in the context of mating competition. We investigated how the presence of females affected salivary cortisol and testosterone levels in males from a small captive group of chimpanzees (Pan troglodytes). Specifically, we assessed whether the presence of females resulted in a rapid increase in salivary cortisol and testosterone levels in the alpha male. We compared the social behavior and salivary hormone concentrations of four males before and after the presentation of receptive females. Three times a day, we collected saliva samples, a useful matrix for investigating short-term hormonal changes, and measured cortisol and testosterone concentration by liquid chromatography-tandem mass spectrometry (LC-MS/MS). The frequency of inter-male aggression increased in the presence of females, indicating intense competition among males. Salivary cortisol levels increased in all males in the presence of females; however, the increase was significantly more pronounced in the alpha male. We found a complex three-way interaction among the presence of females, sampling timings, and male dominance rank in the analysis of salivary testosterone. Contrary to our prediction, a post hoc analysis revealed that salivary testosterone levels decreased after female introduction and that the alpha male did not show a higher level of salivary testosterone. Our study provides experimental evidence suggesting that the presence of females plays a significant role in the rank-related variation in the cortisol levels in male chimpanzees. Furthermore, our findings demonstrate the usefulness of salivary hormones for detecting short-term physiological changes in studies of socioendocrinology.

Captive chimpanzee foraging in a social setting: a test of problem solving, flexibility, and spatial discounting.

In the wild, primates are selective over the routes that they take when foraging and seek out preferred or ephemeral food. Given this, we tested how a group of captive chimpanzees weighed the relative benefits and costs of foraging for food in their environment when a less-preferred food could be obtained with less effort than a more-preferred food. In this study, a social group of six zoo-housed chimpanzees (Pan troglodytes) could collect PVC tokens and exchange them with researchers for food rewards at one of two locations. Food preference tests had revealed that, for these chimpanzees, grapes were a highly-preferred food while carrot pieces were a less-preferred food. The chimpanzees were tested in three phases, each comprised of 30 thirty-minute sessions. In phases 1 and 3, if the chimpanzees exchanged a token at the location they collected them they received a carrot piece (no travel) or they could travel ≥10 m to exchange tokens for grapes at a second location. In phase 2, the chimpanzees had to travel for both rewards (≥10 m for carrot pieces, ≥15 m for grapes). The chimpanzees learned how to exchange tokens for food rewards, but there was individual variation in the time it took for them to make their first exchange and to discover the different exchange locations. Once all the chimpanzees were proficient at exchanging tokens, they exchanged more tokens for grapes (phase 3). However, when travel was required for both rewards (phase 2), the chimpanzees were less likely to work for either reward. Aside from the alpha male, all chimpanzees exchanged tokens for both reward types, demonstrating their ability to explore the available options. Contrary to our predictions, low-ranked individuals made more exchanges than high-ranked individuals, most likely because, in this protocol, chimpanzees could not monopolize the tokens or access to exchange locations. Although the chimpanzees showed a preference for exchanging tokens for their more-preferred food, they appeared to develop strategies to reduce the cost associated with obtaining the grapes, including scrounging rewards and tokens from group mates and carrying more than one token when travelling to the farther exchange location. By testing the chimpanzees in their social group we were able to tease apart the social and individual influences on their decision making and the interplay with the physical demands of the task, which revealed that the chimpanzees were willing to travel farther for better.

Chimpanzees copy dominant and knowledgeable individuals: implications for cultural diversity

Evolutionary theory predicts that natural selection will fashion cognitive biases to guide when, and from whom, individuals acquire social information, but the precise nature of these biases, especially in ecologically valid group contexts, remains unknown. We exposed four captive groups of chimpanzees (Pan troglodytes) to a novel extractive foraging device and, by fitting statistical models, isolated four simultaneously operating transmission biases. These include biases to copy (i) higher-ranking and (ii) expert individuals, and to copy others when (iii) uncertain or (iv) of low rank. High-ranking individuals were relatively un-strategic in their use of acquired knowledge, which, combined with the bias for others to observe them, may explain reports that high innovation rates (in juveniles and subordinates) do not generate a correspondingly high frequency of traditions in chimpanzees. Given the typically low rank of immigrants in chimpanzees, a ‘copying dominants’ bias may contribute to the observed maintenance of distinct cultural repertoires in neighboring communities despite sharing similar ecology and knowledgeable migrants. Thus, a copying dominants strategy may, as often proposed for conformist transmission, and perhaps in concert with it, restrict the accumulation of traditions within chimpanzee communities whilst maintaining cultural diversity.

Preliminary Insights into the Impact of Dietary Starch on the Ciliate, Neobalantidium coli, in Captive Chimpanzees

Infections caused by the intestinal ciliate Neobalantidium coli are asymptomatic in most hosts. In humans and captive African great apes clinical infections occasionally occur, manifested mainly by dysentery; however, factors responsible for development of clinical balantidiasis have not been fully clarified. We studied the effect of dietary starch on the intensities of infection by N. coli in two groups of captive chimpanzees. Adult chimpanzees infected by N. coli from the Hodonín Zoo and from the Brno Zoo, Czech Republic, were fed with a high starch diet (HSD) (average 14.7% of starch) for 14 days, followed by a five-day transition period and subsequently with a period of low starch diet (LoSD) (average 0.1% of starch) for another 14 days. We collected fecal samples during the last seven days of HSD and LoSD and fixed them in 10% formalin. We quantified trophozoites of N. coli using the FLOTAC method. The numbers of N. coli trophozoites were higher during the HSD (mean ± SD: 49.0±134.7) than during the LoSD (3.5±6.8). A generalized linear mixed-effects model revealed significantly lower numbers of the N. coli trophozoites in the feces during the LoSD period in comparison to the HSD period (treatment contrast LoSD vs. HSD: 2.7±0.06 (SE), z = 47.7; p<<0.001). We conclude that our data provide a first indication that starch-rich diet might be responsible for high intensities of infection of N. coli in captive individuals and might predispose them for clinically manifested balantidiasis. We discuss the potential nutritional modifications to host diets that can be implemented in part to control N. coli infections.

Perseverance and food sharing among closely affiliated female chimpanzees

Chimpanzees (Pan troglodytes) have been frequently observed to share food with one another, with numerous hypotheses proposed to explain why. These often focus on reciprocity exchanges for social benefits (e.g., food for grooming, food for sex, affiliation, kinship, and dominance rank) as well as sharing based on begging and deterring harassment. Although previous studies have shown that each of these hypotheses has a viable basis, they have only examined situations in which males have preferential access to food whereby females are required to obtain the food from males. For example, studies on male chimpanzee food sharing take advantage of successful crop-raids and/or acquisitions of meat from hunting, situations that only leave females access to food controlled by male food possessors. This begs the question how and with whom might a female chimpanzee in sole possession of a high-quality food item choose to share? In two large captive groups of chimpanzees, we examined each of the hypotheses with female food possessors of a high-quality food item and compared these data to a previous study examining food transfers from male chimpanzees. Our results show that alpha females shared significantly more with closely affiliated females displaying perseverance, while kinship and dominance rank had no effect. This positive interaction between long-term affiliation and perseverance shows that individuals with whom the female possessor was significantly affiliated received more food while persevering more than those with neutral or avoidant relationships towards her. Furthermore, females with avoidant relationships persevered far less than others, suggesting that this strategy is not equally available to all individuals. In comparison to the mixed-sex trials, females chose to co-feed with other females more than was observed when the alpha male was sharing food. This research indicates that male and female chimpanzees (as possessors of a desired food item) share food in ways influenced by different factors and strategies.

Chimpanzees use multiple strategies to limit aggression and stress during spatial density changes

The regulation of aggression in captive animals is an important welfare concern. Captive environments typically provide limited space for animals and many species exhibit heightened aggression in response to spatial restriction. However, primates appear to regulate aggression under these conditions. These findings have led to the proposal of three models for responses to spatial density changes: the density-intensity, tension-reduction and conflict-avoidance models. Our study aimed to investigate whether spatial restriction in two groups of captive chimpanzees at the Johannesburg Zoo, South Africa, supported the hypotheses of one or more of these models. In addition, a forth model based on the coping hypothesis of stereotypic behaviour was tested. Behavioural observations of both chimpanzee groups were conducted during the nine month reconstruction of the chimpanzee exhibit, and the associated variation in spatial limitation. Both chimpanzee groups used a tension-reduction tactic to limit aggression in the outdoor environments under high spatial density. In the indoor environments, the one (orphan) group of chimpanzees adopted a tension-reduction tactic to limit aggression while the other (family) group adopted a conflict-avoidance tactic. In both groups, indoor high-density conditions generated significant increases in abnormal behaviour. Our findings provide mixed support for the tension-reduction and conflict-avoidance models, while offering no clear evidence for the density-intensity model. The outcomes suggest that the chimpanzees may also have utilised abnormal behaviour as an outlet for the stress of spatial restriction. Together with evidence from other studies, our results suggest that chimpanzees are flexible in their response to the stress of spatial restriction and may employ aggression-mediation tactics in a context dependent manner. With regard to primate welfare, our findings suggest that aggression during spatial restriction may not necessarily be as prominent a welfare concern as previously thought but redirected and abnormal behaviour may still compromise animal wellbeing under spatial restriction.

Chimpanzees share food for many reasons: the role of kinship, reciprocity, social bonds and harassment on food transfers

There is currently great interest in the phylogenetic origins of altruistic behaviour within the primate order. Considerable attention has been focused on chimpanzees, Pan troglodytes, because they are our closest living relatives and participate in a wide range of collective activities, including hunting and food sharing. Food sharing is of particular importance because it plays a critical role in the human foraging niche, but food sharing among adults is rare in nonhuman primates. Some research suggests that chimpanzees selectively share meat with reciprocating partners and allies, while other work indicates that chimpanzees primarily share to reduce harassment from other group members (tolerated theft). We examined the effects of kinship, relationship quality, reciprocity and the intensity of solicitations on the pattern of food transfers in six captive groups of chimpanzees. We observed events that occurred after the chimpanzees were provisioned with large frozen juice disks. These disks share some properties with prey carcasses: they are a valued, but limited, resource; they take a considerable period of time to consume; they can be monopolized by one individual, but bits can be broken off and transferred to others. Our analyses suggest that food transfers serve multiple functions for chimpanzees. Individuals may use food transfers to enhance the welfare of closely related group members, strengthen social relationships with favoured partners and reduce the costs of persistent solicitations.

Shade as a thermoregulatory resource for captive chimpanzees

In nature, the thermoregulatory strategies of species have evolved in response to the environmental conditions in which they live. Primates display extensive behavioural flexibility but few have examined the role of this behavioural flexibility with regard to thermoregulation. Chimpanzees, under free-living conditions, utilise natural vertical light/thermal gradients in rainforest canopies as a means of thermoregulation, moving along the gradient to maintain homoeostasis. In contrast, captive chimpanzees are often housed at latitudes outside their natural range under conditions that do not allow for this natural behavioural thermoregulation. We investigated the use of shade by captive chimpanzees as a behavioural thermoregulatory strategy at the Johannesburg Zoo, South Africa. We performed behavioural observations over the austral winter/spring period on a group of captive chimpanzees recording their behaviour in relation to shade use. Despite experiencing temperatures in Johannesburg below or just within their thermoneutral zone, chimpanzees consistently spent more time in shade than in direct sun. This pattern of shade use was most pronounced for the hotter midday period than for other times, was not dependent on the overall availability of shade within the enclosure and was not predicted by mean daily or hourly temperatures nor thermal maxima or minima. Instead, ultraviolet radiation and humidity levels appear to predict the observed patterns of shade utilisation and these findings suggest that chimpanzees in captivity adopt a sun-avoidance strategy, possibly as a result of the rapid heat gain associated with their dark skin and pelage. The findings suggest that chimpanzees are flexible when responding to the thermal challenges associated with housing outside of their natural environment.

Volunteer experience influences the conclusions of behavioural experiments

Volunteers offer an inexpensive and rapid means of collecting behavioural data, but their reliability is often overlooked. Past research has suggested that observers that were inexperienced are equally adept at recording behaviour as experienced observers, and inexperience was regarded as being merely unfamiliar with a sampling technique but not unknowledgeable about behaviour. The aims of our study were (i) to investigate the reliability of relatively naïve volunteers (i.e. those with no prior behavioural scoring experience) as behavioural data collectors; and (ii) to test the influence of the strength of inter-observer concordance on the outcome of testing a specific ethological hypothesis. Two cohorts of volunteers (high school and university students) conducted observations on a group of captive chimpanzees, simultaneously with an experienced observer (LD), recording behaviour and the location of the chimpanzees in their enclosure. Kendall's Tau agreement scores and odds ratios indicated poor agreement between inexperienced volunteers and the experienced observer, regardless of the educational experience of the volunteers and difficulty of the behaviour scored. We compared the data between the volunteers and experienced observer by independently testing each dataset with regard to the hypothesis that the chimpanzees were stressed by being in close proximity to the public. The school cohort data supported the hypothesis, while the time-matched experienced observer data suggested no relationship between public proximity and stress in chimpanzees. A separate analysis of the university cohort and time-matched experienced observer data both indicated that chimpanzees were more stressed at locations away from the public. These findings suggest that inter-observer agreement scores offer insights into the precision of data but not accuracy. Furthermore, the use of volunteers as data collectors should be assessed in relation to the aims of the study in question, since volunteers may be appropriate for studies of general patterns of behaviour but not for detailed ethological examinations.

Assessing the effects of cognitive experiments on the welfare of captive chimpanzees (Pan troglodytes) by direct comparison of activity budget between wild and captive chimpanzees

We investigated the effects of cognitive experiments by direct comparison of activity budgets between wild and captive chimpanzees. One goal of captive management is to ensure that the activity budgets of captive animals are as similar as possible to those of their wild counterparts. However, such similarity has rarely been achieved. We compared the activity budget among three groups of chimpanzees: wild chimpanzees in Bossou (Guinea, n = 10), and captive chimpanzees who participated in cognitive experiments (experimental chimpanzees, n = 6) or did not participate in the experiments (nonexperimental chimpanzees, n = 6) at the Primate Research Institute (Japan). The experimental chimpanzees voluntarily participated in computer-controlled cognitive tasks and small pieces of fruits were provided as rewards. The data from captivity were obtained on the experimental days (weekdays) and nonexperimental days (weekends). In both study sites, we followed each chimpanzee from about 7 a.m. until the time when chimpanzees started to rest in the evening. The behaviors were recorded every 1 min. The results showed that on weekdays, feeding time and resting time of the experimental chimpanzees were almost the same as those of wild chimpanzees. However, for the nonexperimental chimpanzees, feeding time was significantly shorter and resting time was longer than those of the wild chimpanzees. In contrast, no difference was found in feeding time or resting time of the two groups of captive chimpanzees on weekends. The results suggested that the cognitive experiments worked as an efficient method for food-based enrichment.

Individual Differences in Long-term Cognitive Testing in a Group of Captive Chimpanzees

Seven chimpanzees had participated in cognitive tasks from the time they were approximately 18 months to approximately 16 years of age when the data presented here was analyzed. Testing covered a wide range of tasks, which we categorized broadly as measuring their understanding of aspects of either their social or physical environments. Therefore, we could test whether individuals who excelled on ‘social’ tasks, also excelled on ‘physical’ tests. We also categorized our measures as ones of acquisition, criterion, retention or transfer of skill. Thus, we could determine whether individuals who mastered tasks quickly were also those who performed, remembered and generalized tasks most accurately. We were interested in whether there were consistent patterns in cognitive skills across tasks and measures. Results of our analyses indicate that, as with humans, chimpanzees vary in their performance across some measures, although some differences in cognitive skill between individuals are also consistent across measures and tasks. The results have implications for questions concerning domain generality or specificity of cognitive skills in another primate species.//

Grooming network cohesion and the role of individuals in a captive chimpanzee group

Social network analysis offers new tools to study the social structure of primate groups. We used social network analysis to investigate the cohesiveness of a grooming network in a captive chimpanzee group (N = 17) and the role that individuals may play in it. Using data from a year-long observation, we constructed an unweighted social network of preferred grooming interactions by retaining only those dyads that groomed above the group mean. This choice of criterion was validated by the finding that the properties of the unweighted network correlated with the properties of a weighted network (i.e. a network representing the frequency of grooming interactions) constructed from the same data. To investigate group cohesion, we tested the resilience of the unweighted grooming network to the removal of central individuals (i.e. individuals with high betweenness centrality). The network fragmented more after the removal of individuals with high betweenness centrality than after the removal of random individuals. Central individuals played a pivotal role in maintaining the network's cohesiveness, and we suggest that this may be a typical property of affiliative networks like grooming networks. We found that the grooming network correlated with kinship and age, and that individuals with higher social status occupied more central positions in the network. Overall, the grooming network showed a heterogeneous structure, yet did not exhibit scale-free properties similar to many other primate networks. We discuss our results in light of recent findings on animal social networks and chimpanzee grooming.

Third‐party postconflict affiliation of aggressors in chimpanzees

Postconflict management strategies have been defined as any postconflict interaction that mitigates the negative consequences of the preceding agonistic conflict. Although most studies have investigated postconflict interactions between former opponents or between victims and uninvolved bystanders, interactions between aggressors and bystanders have received much less attention. In this study, we examined a database of 1,102 agonistic interactions and their corresponding postconflict periods in two outdoor-housed groups of captive chimpanzees in order to test the occurrence of postconflict third-party affiliation of aggressors. Our results confirmed the occurrence of appeasement, i.e. postconflict affiliation by a bystander toward an aggressor, but failed to detect the occurrence of postconflict affiliation directed from aggressors toward bystanders. Appeasement rates did not differ according to the sex of the involved individuals. In addition, appeasement occurred more often in the absence of reconciliation than after its occurrence suggesting that appeasement may act as an alternative to reconciliation when the latter fails to occur. Both study groups showed behavioral specificity for appeasement, i.e. context-specific use of certain behaviors, supporting the view that chimpanzees exhibit highly visible explicit postconflict affiliation.

Tool use in a group of captive chimpanzees I. Escape.

A group of 4 female and 3 male adolescent wild-born chimpanzees was observed for 7 months in an outdoor enclosure. Use of tools aimed at escaping from this enclosure is decribed, the two main techniques being the use of poles as ladders and sticks as pitons.