Post-conflict affiliation by chimpanzees with aggressors: other-oriented versus selfish political strategy

Consolation, i.e., post-conflict affiliation directed from bystanders to recent victims of aggression, has recently acquired an important role in the debate about empathy in great apes. Although similar contacts have been also described for aggressors, i.e., appeasement, they have received far less attention and their function and underlying mechanisms remain largely unknown. An exceptionally large database of spontaneous conflict and post-conflict interactions in two outdoor-housed groups of chimpanzees lends support to the notion that affiliation toward aggressors reduces the latter's aggressive tendencies in that further aggression was less frequent after the occurrence of the affiliation. However, bystander affiliation toward aggressors occurred disproportionally between individuals that were socially close (i.e., affiliation partners) which suggest that it did not function to protect the actor itself against redirected aggression. Contrary to consolation behavior, it was provided most often by adult males and directed toward high ranking males, whereas females engaged less often in this behavior both as actors and recipients, suggesting that affiliation with aggressors is unlikely to be a reaction to the distress of others. We propose that bystander affiliation toward aggressors may function to strengthen bonds between valuable partners, probably as part of political strategies. Our findings also suggest that this post-conflict behavior may act as an alternative to reconciliation, i.e., post-conflict affiliation between opponents, in that it is more common when opponents fail to reconcile.     

Reconciliation, consolation and postconflict behavioral specificity in chimpanzees

Conflicts of interest arise regularly in the lives of all group-living animals and may escalate into aggressive conflicts. The costs of aggressive escalation can be reduced through peaceful postconflict interactions. This study investigated the postconflict behavior of 22 adult chimpanzees at Chester Zoo. The occurrence of reconciliation, i.e. the postconflict affiliative reunion between conflict opponents, and consolation, i.e. a postconflict affiliative interaction directed from a third party to the recipient of aggression, were demonstrated. Consolation was more likely to occur in the absence of reconciliation than after reconciliation, and reconciliation was more likely to occur in the absence of consolation than after consolation, supporting the hypothesis that consolation acts as a substitute for reconciliation when the latter fails to occur. Evidence for behavioral specificity, i.e. context-specific use of certain behaviors, was found for both reconciliation and consolation, which, along with high conciliatory tendencies, suggests an explicit style of postconflict behavior in the study subjects.     

The Occurrence of Postconflict Skills in Captive Immature Chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Conflict management strategies can reduce costs of aggressive competition in group-living animals. Postconflict behaviors such as reconciliation and third-party postconflict affiliation are widely accepted as social skills in primates and have been demonstrated in many species. Although immature primates possess a repertoire of species-specific behaviors, it is thought that they gradually develop appropriate social skills throughout prolonged juvenility to establish and maintain complex social relationships within their group. We examined the occurrence of postconflict skills in five immature chimpanzees (Pan troglodytes) over 15 mo, focusing on interactions that were not with the subject’s mother. We observed reconciliation, with conciliatory tendencies comparable to adults, and provide the first evidence that captive immature chimpanzees commonly reconciled using social play. However, immatures were not more likely to reconcile valuable than nonvaluable relationships. We also observed third party postconflict affiliation although at a lower level than reported for adults. Our results provide evidence for postconflict skills in immature chimpanzees but the lack of higher conciliatory tendency with valuable partners and low occurrence of third-party affiliation indicates extended juvenility may be required refine these skills. Further work is needed to investigate whether these behaviors have the same function and effectiveness as those found in adults.     

Postconflict Behavior of Spectacled Leaf Monkeys (Trachypithecus obscurus). I. Reconciliation

Studies of postconflict behavior have focused on Old World primates, particularly on members of the cercopithecines and chimpanzees. We present data on patterns of reconciliation in two captive groups of spectacled leaf monkeys, members of the Colobinae. Reconciliation occurred at a high rate relative to that in other primate species and within the first minute following a conflict. Friendly postconflict contact was usually initiated by aggressors in one group, and by recipients in the other. But, there was no difference in postconflict initiative to contact in comparison with MCs. These two groups showed behavioral specificity in the context of postconflict reunions. Ventroventral embracing was a common, explicit (de Waal, 1993) form of reconciliation. The intensity of aggression and situations in which the victim responded aggressively had no effect on the likelihood of reconciliation occurring. The postconflict behavior of both groups lends further support to the relationship-quality hypothesis (Aureli et al., 1989). Kinship and rank distance and had no effect on reconciliation, though affiliation levels correlated positively with the tendency to reconcile. These results indicate that spectacled langurs are among the more tolerant of the species studied so far.     

Postconflict Affiliation of Aggressors in <Emphasis Type="Italic">Papio hamadryas</Emphasis>

Researchers have associated variation in the occurrence of postconflict behaviors with variation in the relationship quality between involved individuals. Apart from those on the great apes, the vast majority of postconflict studies involved female-bonded species and focused on the victim. We examined postconflict affiliation involving the aggressor in a non-female-bonded species, hamadryas baboons (Papio hamadryas), and investigated the influence of relationship quality on the occurrence of 2 types of postconflict affiliative behaviors. Hamadryas baboons reconciled after a conflict and the highest rate of reconciliation was between one-male unit (OMU) leader males and their females. Via direct measures of relationship characteristics we also found that partners with higher-quality relationship, e.g., highly affiliative dyads and allies, showed higher levels of conciliatory tendency than dyads with lower-quality relationship, e.g., less affiliative dyads and non-allies. We found evidence of postconflict third-party affiliation initiated by aggressors, but not by third parties. Further, aggressor-initiated affiliation was more likely with individuals of the same OMU and individuals with which the aggressor maintained a strongly affiliative relationship. Our findings provide further support for relationship quality as a fundamental underlying factor not only in reconciliation, but also in postconflict affiliations involving third parties.     

Functions of post‐conflict affiliation with a bystander differ between aggressors and victims in Japanese macaques

Post‐conflict affiliative interactions with a bystander, an uninvolved third party, have been reported in several species. The functions of these interactions could vary depending on whether the individual was an aggressor or a victim in a conflict. We examined the occurrence of and three potential functions for bystander affiliation—substitute for reconciliation, self‐protection and consolation (stress reduction)—in a free‐ranging group of Japanese macaques. We predicted that in this despotic species, the function of bystander affiliations for victims was self‐protection, whereas this was not so for aggressors. We found that compared to control situations, both aggressors and victims engaged in bystander affiliation soon after a conflict, confirming the occurrence of bystander affiliation. Victims were less likely to receive further aggression when bystander affiliation occurred than when no such interaction occurred, consistent with the explanation that post‐conflict bystander affiliation functions as self‐protection for victims. Aggressors were less likely to perform self‐scratching, an indicator of post‐conflict stress, when bystander affiliation occurred. This suggests that such affiliations benefited aggressors by reducing stress. Self‐scratching was not less frequent after affiliation with a bystander who was familiar with the former opponent, indicating that substitute reconciliation did not apply for either aggressors or victims. When bystander affiliation occurred, the bystander was a more familiar individual for aggressors and victims than the former opponent was. In conclusion, aggressors benefited in stress reduction, whereas victims benefited in self‐protection through affiliation with bystanders with whom they have a good relationship. Our study indicated that monkeys cope with post‐conflict problems according to their roles or situations not only by reconciliation but also by bystander affiliation.     

Affiliation Between Aggressors and Third Parties Following Conflicts in Long-Tailed Macaques (Macaca fascicularis)

Studies on cercopithecine monkeys have shown that soon after an agonistic conflict, victims have increased rates of affiliation with the agressor—reconciliation—but not with other group members. Postconflict affiliation is thought to function to restore disturbed relationships and to reduce social tension. This study on a captive group of long-tailed macaques (Macaca fascicularis) is focused on postconflict affiliative behavior of the aggressor. Increased rates of contact between female aggressors and kin of the victim occurred, as well as between female aggressors and their own kin. Furthermore, there were increased rates of contact between aggressors—males and females—and other group members. The increase in contacts with the victim's kin was selective, i.e., it could not be ascribed to the increased contact tendency with group members in general, and was not a side effect of the aggressor's proximity to the victim due to reconciliation. The increase in contacts with own kin was not selective. The fact that male aggressors do not have increased postconflict contacts with their kin or with kin of the victim is in agreement with the notion that males are less integrated in the nepotistic matrilineal network than females are. The fact that studies by others that focused on the victim evidence no increase in postconflict contacts with kin of the opponent or with other group members may be explained by the aggressor's larger influence over the postconflict situation: to reduce social tension, it might be more effective to affiliate with the aggressor than with the victim. Our findings emphasize that conflicts influence the behavior of other monkeys besides the direct contestants and, thus, indicate that the disturbance of social homeostasis is a matter of concern for all group members.     

Receiving post-conflict affiliation from the enemy's friend reconciles former opponents

The adaptive function of bystander initiated post-conflict affiliation (also: consolation & appeasement) has been debated for 30 years. Three influential hypotheses compete for the most likely explanation but have not previously been tested with a single data set. The consolation hypothesis argues that bystander affiliation calms the victim and reduces their stress levels. The self-protection hypothesis proposes that a bystander offers affiliation to either opponent to protect himself from redirected aggression by this individual. The relationship-repair hypothesis suggests a bystander can substitute for a friend to reconcile the friend with the friend''s former opponent. Here, we contrast all three hypotheses and tested their predictions with data on wild chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d''Ivoire. We examined the first and second post-conflict interactions with respect to both the dyadic and triadic relationships between the bystander and the two opponents. Results showed that female bystanders offered affiliation to their aggressor friends and the victims of their friends, while male bystanders offered affiliation to their victim friends and the aggressors of their friends. For both sexes, bystander affiliation resulted in a subsequent interaction pattern that is expected for direct reconciliation. Bystander affiliation offered to the opponent''s friend was more likely to lead to affiliation among opponents in their subsequent interaction. Also, tolerance levels among former opponents were reset to normal levels. In conclusion, this study provides strong evidence for the relationship-repair hypothesis, moderate evidence for the consolation hypothesis and no evidence for the self-protection hypothesis. Furthermore, that bystanders can repair a relationship on behalf of their friend indicates that recipient chimpanzees are aware of the relationships between others, even when they are not kin. This presents a mechanism through which chimpanzees may gain benefits from social knowledge.     

Postconflict Affiliation and Stress-Related Behavior of Long-Tailed Macaque Aggressors

Previous studies on macaques and baboons showed that after agonistic conflicts aggressees as well as aggressors show an increase in stress-related behavior such as scratching. Reconciliation reduces stress-related behavior of the aggressee. We investigated the influence of various affiliative postconflict behaviors of the aggressor on the aggressor's scratching rates in captive long-tailed macaques: reconciliation, contacts with the aggressee's kin (or substitute reconciliation), and contact with other group members (or triadic affiliation). After a conflict, the aggressor showed an increase in rates of scratching. Scratching rates were reduced after reconciled conflicts compared to nonreconciled conflicts. Substitute reconciliation did not reduce scratching when we controlled for the influence of reconciliation, i.e., the aggressor might not interpret it as a substitute for reconciliation. Triadic affiliation did not reduce scratching rates, hence, triadic affiliation probably does not console the aggressor. Scratching rates after reconciliation are significantly lower than scratching rates after triadic affiliation. This proves that the stress-reducing effect of reconciliation is not due to the calming effect of general body contact but that the stress reduction is specifically associated with contacts with the former opponent. The contestants are anxious about their relationship, and only reconciliation takes away this anxiety. Reconciliation is thus an important social repair strategy.     

Postconflict Affiliation Between Former Opponents in Macaca thibetana on Mt. Huangshan, China

We describe basic patterns of postconflict affiliation between former opponents within a group of wild, provisioned Tibetan macaques Macaca thibetana on Mt. Huangshan, China. Like most primates studied to date, Tibetan macaques reconciled, i.e., overall they engaged in affiliative interaction with opponents at higher rates immediately after an aggressive conflict than at other times. Probabilities of affiliation were enhanced ≤30 s after the end of hostilities. However when we examined sex partner combinations separately, we found unequivocal evidence for reconciliation only for male-male dyads. Tolerant interaction among other partner combinations apparently was not disrupted after a conflict, presumably obviating the need to reconcile. One aspect of reconciliation among males was consistent with other indications of a despotic dominance style: aggressors initiated a higher proportion of affiliative interactions after a conflict than at other times. Another aspect of reconciliation was more typical of relaxed dominance styles: males used specialized behaviors (embraces and same-sex mounts) disproportionately to reconcile. We also found inconsistent evidence for the valuable relationship hypothesis; probabilities of reconciliation were enhanced for male-male dyads with the closest affiliative relationships, but not for those that displayed the most tolerance or mutual agonistic support. We discuss reconciliation and other aspects of conflict management among males in the context of a group with nearly even sex ratios and high male-male mating competition.     

The Function and Determinants of Reconciliation in <Emphasis Type="BoldItalic">Pan troglodytes</Emphasis>

Reconciliation (the postconflict affiliative reunion between former opponents) may mitigate costs of aggressive conflict by repairing the opponents’ relationship and reducing stress. We showed that postconflict levels of self-directed behavior were lower after reconciliation than when reconciliation did not occur in chimpanzees (Pan troglodytes) at Chester Zoo, providing support for a stress-alleviating function for reconciliation. Further, we investigated the effects of multiple factors on the occurrence of reconciliation using generalized linear mixed models. We performed 2 separate analyses, a “traditional” analysis and a “targeted” analysis. The former included variables previously used to assess the occurrence of reconciliation in primates, i.e., conflict characteristics, sex combination, and a simple measure of relationship value. The latter included species-specific variables such as the occurrence of consolation (postconflict affiliation from a bystander to the recipient of aggression); initiation of the conflict with a bluff display; and measures of relationship value, compatibility, and security specific to the study group. Whereas the traditional analysis showed that female-female dyads and valuable partners were most likely to reconcile, the targeted analysis showed that reconciliation was less likely to occur when consolation took place or when aggression was initiated with bluff displays. Further analyses revealed that the effect of sex-combination on reconciliation was due to its intercorrelation with bluff display. This study highlights the importance of considering variables specific to the study species and group when investigating the determinants of reconciliation and warns against premature interpretation of results without due consideration for all other possible determinants.     

Conflict and postconflict behaviour in captive black-and-white snub-nosed monkeys (<Emphasis Type="Italic">Rhinopithecus bieti</Emphasis>)

Black-and-white snub-nosed monkeys (Rhinopithecus bieti) have almost never been the subject of any behavioural observations in captivity. This study was aimed at providing preliminary information about agonistic and reconciliation behaviour in a group kept at the Kunming Institute of Zoology in China. Established procedures were used for this investigation (i.e., the postconflict/matched-control method and the time-rule method). Intra-group aggression rates were quite low. Postconflict affiliation as well as selective attraction of former opponents to each other following conflicts was demonstrated. Former opponents contacted each other earlier in postconflict periods than in matched-control periods. The average conciliatory tendency of all focal individuals combined was 54.5%. After an agonistic interaction, the first affiliative contact between former aggressors usually took place within the first minute. The behaviours most often shown as first affiliations after a conflict were body contact, mount, touch, and hold-lumbar, of which the latter is an explicit reconciliatory gesture. Furthermore, the adult male intervened non-aggressively in 84% of all conflicts (n=25) among the adult females. Overall, the patterns of aggression and reconciliation observed in R. bieti bear many of the traits that characterise tolerant primate species.     

Postconflict affiliative contacts between former opponents among wild moor macaques (Macaca maurus)

Agonistic interactions and postconflict behavior of moor macaques (Macaca maurus) were studied in their natural habitat, South Sulawesi, Indonesia. Individuals were involved in 0.28 agonistic interactions per hour. Forty-two percent of agonistic interactions were followed by affiliative contacts between the former opponents. Such affiliative contacts occurred within 2 minutes of the agonistic interaction and were most often initiated by the victim's approach to the aggressor. The most common postconflict behavior was grooming of the aggressor by the victim. The present study provides further evidence of a positive correlation between a high tendency for postconflict affiliative contact and a relaxed dominance style. Captivity might not alter the expression of postconflict behavior. The present study suggests that it is possible to demonstrate the occurrence of reconciliation a posteriori by using data obtained by general focal observation protocols. © 1996 Wiley-Liss, Inc.     

Testing the function of reconciliation and third-party affiliation for aggressors in hamadryas baboons (Papio hamadryas hamadryas)

In social groups, agonistic conflicts can have different negative consequences. Several post-conflict interactions have been suggested as post-conflict management behaviors to mitigate those negative effects. In this study, we investigated the function of two post-conflict behaviors--reconciliation and aggressor-initiated third-party affiliation--on the aggressor's levels of post-conflict anxiety and aggression in a large colony of hamadryas baboons. We also examined variation in the aggressor's levels of post-conflict anxiety as a function of relationship quality between the opponents as predicted by the Integrated Hypothesis. We found that after conflicts hamadryas baboon aggressors showed increased rates of anxiety-related behaviors and that they were also more likely to be involved in renewed aggressive interactions. Although both reconciliation and aggressor-initiated third-party affiliation reduced the probability of receiving post-conflict aggression, only reconciliation reduced the rates of anxiety-related behaviors, suggesting that the aggressors' post-conflict anxiety might be owing mainly to the damage that the conflict causes to their relationship with the victim. Furthermore, aggressor's rates of post-conflict anxiety were higher after conflicts with individuals with whom they had a high-quality relationship, supporting the idea that levels of post-conflict anxiety mediate the occurrence of reconciliation depending on the quality of the relationship with former opponent as predicted by the Integrated Hypothesis.     

Postconflict Behavior of Spectacled Leaf Monkeys (Trachypithecus obscurus). II. Contact with Third Parties

We studied the postconflict interactions with group members other than the former opponent in two groups of spectacled leaf monkeys. We found no evidence of redirection of aggression towards other group members. Victims and aggressors sought affiliative contacts with uninvolved third parties. There was no evidence for consolation—affiliative contact initiated by an uninvolved third party, directed towards victims of aggression—in either group when all affiliative behaviors were considered. However, embracing was a characteristic first-contact interaction between individuals involved in aggression and third parties. This finding mirrors the results concerning reconciliation in spectacled leaf monkeys. Accordingly, embracing may be a true consolatory behavior in this species. When contacts with third parties occurred before or in the absence of reconciliation, the timing of such contacts fell within the time window during which reconciliation normally occurs. These contacts also resulted in affiliation levels twice that of baseline levels, supporting the idea that these contacts may function as a form of substitute reconciliation. We discuss these results in light of recent theories concerning postconflict behavior in primates.     

Chimpanzees Extract Social Information from Agonistic Screams

Chimpanzee (Pan troglodytes) agonistic screams are graded vocal signals that are produced in a context-specific manner. Screams given by aggressors and victims can be discriminated based on their acoustic structure but the mechanisms of listener comprehension of these calls are currently unknown. In this study, we show that chimpanzees extract social information from these vocal signals that, combined with their more general social knowledge, enables them to understand the nature of out-of-sight social interactions. In playback experiments, we broadcast congruent and incongruent sequences of agonistic calls and monitored the response of bystanders. Congruent sequences were in accordance with existing social dominance relations; incongruent ones violated them. Subjects looked significantly longer at incongruent sequences, despite them being acoustically less salient (fewer call types from fewer individuals) than congruent ones. We concluded that chimpanzees categorised an apparently simple acoustic signal into victim and aggressor screams and used pragmatics to form inferences about third-party interactions they could not see.     

The Importance of Considering the Behavioral Form of Reconciliation in Studies of Conflict Resolution

Reconciliation is the most extensively studied conflict resolution mechanism in animal societies. However, despite the extensive literature on this topic, behaviors considered to represent postconflict affiliation have not been consistent across studies of reconciliation. Critically, reconciliation is usually defined as postconflict contact affiliation, e.g., grooming, and the importance of including interopponent distance regulation is often neglected. Moreover, to date, no study has simultaneously investigated different behavioral forms of reconciliation. We tested in two groups of wild Barbary macaques (Macaca sylvanus) the relative importance of postconflict close proximity and grooming in the mediation of two important costs of aggression: damage to the opponent’s social relationship and elevated postconflict anxiety. We provide evidence that close-proximity approaches function to resolve conflicts: Close-proximity approaches reduced the victim’s postconflict anxiety and were predicted by the quality of the social relationship with the opponent. Moreover, postconflict grooming alone, although predicted by the quality of the opponent’s social relationship, did not influence the victim’s elevated postconflict anxiety. Our results suggest that interopponent distance regulation plays an important role in reconciling the costs of aggression in Barbary macaques. We advocate that further efforts should be made to test which behaviors play a role in conflict resolution in different species. This is important because even closely related species may differ in the function of behaviors that superficially appear to be rather similar. Moreover, the choice of behaviors used to study conflict resolution determines the frequency with which reconciliation is observed and can thus bias comparisons across species.     

Postconflict social interactions among barbary macaques (Macaca sylvanus)

We investigated the occurrence of postconflict social interactions in a group of Barbary macaques (Macaca sylvanus)that live in a large enclosure. We collected data during the mating and the nonmating seasons and analyzed them via two methods. Separate analyses for the two seasons yielded very similar results. Reconciliation and redirection occurred. But we found no evidence for consolation and postconflict kin-oriented affiliation. The consistency of these findings with those on other species of Macacaindicates that reconciliation and redirection are typical postconflict social interactions of macaque victims.     

Reconciliation, relationship quality, and postconflict anxiety: testing the integrated hypothesis in captive chimpanzees

Reconciliation is a conflict resolution mechanism that is common to many gregarious species with individualized societies. Reconciliation repairs the damaged relationship between the opponents and decreases postconflict (PC) anxiety. The "integrated hypothesis" links the quality of the opponents' relationship to PC anxiety, since it proposes that conflicts among partners with high relationship quality will yield high levels of PC anxiety, which in turn will lead to an increased likelihood of reconciliation. We tested the integrated hypothesis in captive chimpanzees (Pan troglodytes) in the Arnhem Zoo, The Netherlands. We applied the standard PC/matched control (MC) method. Our results mostly support the integrated hypothesis, in that more valuable and compatible partners (i.e., males and frequent groomers) reconciled more often than less valuable and weakly compatible partners (i.e., females and infrequent groomers). In addition, PC anxiety was higher after conflicts among males than among females. Emotional arousal thus appears to be a mediator facilitating reconciliation. However, in contrast to the predictions derived from the integrated hypothesis, PC anxiety appeared only in aggressees, and not in aggressors, of conflicts. This suggests that while relationship quality determines PC anxiety, it is dependent on the role of the participants in the conflict.     

Reconciliation in Captive Chimpanzees: A Reevaluation with Controlled Methods

Affiliative postconflict reunions—reconciliations—of former opponents were first demonstrated in the chimpanzees at the Arnhem Zoo. Since then methods have been considerably refined, and reconciliation has been demonstrated in a large number of primates and also some gregarious nonprimates. This study, conducted with a different captive group, is the first to use the revised methodology with chimpanzees. We analyzed a total of 297 agonistic conflicts with the PC–MC method: we observed focal individuals for 15 min after a conflict and during matched control observations the next day. The mean conciliatory tendency of the 16 chimpanzees was 41%, with a range in different age-sex classes of 58% (among adult females) to 19% (among adult vs. immature males). After conflicts, former opponents were selectively attracted to one another. Preferential contact with previous opponents persisted when activity level during matched controls was controlled for statistically. Opponents that were frequent grooming partners reconciled more frequently, but the frequency of agonistic support had no such effect. Our findings thus confirm the existence of reconciliation in chimpanzees, which show one of the highest conciliatory tendencies among primate species.