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.     

Grooming coercion and the post-conflict trading of social services in wild Barbary macaques

In animal and human societies, social services such as protection from predators are often exchanged between group members. The tactics that individuals display to obtain a service depend on its value and on differences between individuals in their capacity to aggressively obtain it. Here we analysed the exchange of valuable social services (i.e. grooming and relationship repair) in the aftermath of a conflict, in wild Barbary macaques (Macaca sylvanus). The relationship repair function of post-conflict affiliation (i.e. reconciliation) was apparent in the victim but not in the aggressor. Conversely, we found evidence for grooming coercion by the aggressor; when the victim failed to give grooming soon after a conflict they received renewed aggression from the aggressor. We argue that post-conflict affiliation between former opponents can be better described as a trading of social services rather than coercion alone, as both animals obtain some benefits (i.e. grooming for the aggressor and relationship repair for the victim). Our study is the first to test the importance of social coercion in the aftermath of a conflict. Differences in competitive abilities can affect the exchange of services and the occurrence of social coercion in animal societies. This may also help explain the variance between populations and species in their social behaviour and conflict management strategies.     

Ravens reconcile after aggressive conflicts with valuable partners

Reconciliation, a post-conflict affiliative interaction between former opponents, is an important mechanism for reducing the costs of aggressive conflict in primates and some other mammals as it may repair the opponents' relationship and reduce post-conflict distress. Opponents who share a valuable relationship are expected to be more likely to reconcile as for such partners the benefits of relationship repair should outweigh the risk of renewed aggression. In birds, however, post-conflict behavior has thus far been marked by an apparent absence of reconciliation, suggested to result either from differing avian and mammalian strategies or because birds may not share valuable relationships with partners with whom they engage in aggressive conflict. Here, we demonstrate the occurrence of reconciliation in a group of captive subadult ravens (Corvus corax) and show that it is more likely to occur after conflicts between partners who share a valuable relationship. Furthermore, former opponents were less likely to engage in renewed aggression following reconciliation, suggesting that reconciliation repairs damage caused to their relationship by the preceding conflict. Our findings suggest not only that primate-like valuable relationships exist outside the pair bond in birds, but that such partners may employ the same mechanisms in birds as in primates to ensure that the benefits afforded by their relationships are maintained even when conflicts of interest escalate into aggression. These results provide further support for a convergent evolution of social strategies in avian and mammalian species.     

Post-conflict Behaviour of Wild Olive Baboons. II. Stress and Self-directed Behaviour

Previous studies indicate that the proximate function of post-conflict affiliative interaction among primates is to reduce a victim's uncertainty about its opponent's future behaviour: the ‘uncertainty reduction hypothesis’ (Aureli & van Schaik 1991; Ethology, 89 , 101–114). This study confirms and extends these results, demonstrating that they are neither a product of captivity nor specific to macaques: both victims and initiators of aggression in a large group of wild olive baboons exhibited elevated rates of self-directed behaviour (SDB) - scratching, autogrooming, body-shaking and yawning - in a 10-min post-conflict period. During this period, they were more likely to receive further aggression. Reconciliation reduced both SDB and the incidence of further aggression. However, reconciliation only reduced SDB among individuals involved in conflicts in which both parties exchanged aggression. It is suggested that aggressors in unilateral conflicts were aroused rather than uncertain and that their victims' lack of control over post-conflict interactions (which tended to be initiated by their opponents) prevented them from benefiting from reconciliation in the same fashion as longtailed macaque victims, which frequently initiated reconciliation.     

Dyadic and triadic reconciliation in pigtail macaques (Macaca nemestrina)

The tendency in primates for former antagonists to approach and affiliate following aggression has been termed reconciliation because the response is thought to resolve social conflicts produced by aggression. In primate societies, however, an aggressive interaction between two individuals often spreads to include other group members, especially the kin of the combatants. If post conflict affiliation resolves aggressive conflicts in a group, then affiliative increases might occur between combatants and the kin of their opponents following aggression as well as between former opponents. This hypothesis was tested in a captive group of 39 pigtail macaques (Macaca nemestrina) by comparing affiliative response frequencies of combatants during the 5 minute period following aggression to affiliative response frequencies during 5 minute baseline periods not preceded by aggressive activity. Following aggression, affiliation rates increased between combatants and their opponents, aggressors and the kin of their opponents, and aggressors and their own kin. Additional analyses showed that aggression among kin was reconciled more often than aggression among nonkin. Recipients of aggression reconciled with their attackers more often than aggressors reconciled with their victims. Animals with similar dominance ranks reconciled proportionately more often than those with large rank disparities and aggressive infractions of a calculated dominance hierarchy were reconciled more often than attacks consistent with the hierarchy. Results suggest that both dyadic and triadic reconciliations occur in M. nemestrina and that compared to other primate species M. nemestrina exhibit a moderate-to-high conciliatory tendency.     

Post-Conflict Affiliation in Two Captive Groups of Black-and-White Guereza Colobus guereza

Post‐conflict affiliation was studied in two captive groups of black‐and‐white guereza (Colobus guereza), a folivorous, arboreal species which lives in harems. The overall level of aggression was low and each individual was involved in only 0.054 aggressive interactions per hour. After 132 aggressive interactions observed during 364 h, former antagonists were found to seek each other out selectively and exchange affiliative behaviours. The first affiliative behaviour was likely to occur within the first minute after the end of the conflict. The post‐conflict affiliative behaviours most often seen were proximity, social grooming, embrace, reciprocal low‐intensity mouth‐opening, groom present and gentle hand touch, in descending order. The pattern of post‐conflict affiliation is thus similar to that observed in most other primate species studied. One notable exception seems to be the relatively high proportion of conflicts after which former opponents continue to remain within arm's reach (in 1/3 of the conflicts). If these instances of remaining‐in‐proximity were excluded, the corrected conciliatory tendency according to Veenema et al. (1994, Behav. Proc. 31, 29–38) was 45% (60% if they were included).     

Rook,But Not Jackdaw,Post‐Conflict Third‐Party Affiliation Reduces Aggression for Aggressors

Post‐conflict (PC) affiliation refers to positive social interactions that occur after fights. Although this behavior has been widely studied, its functions are rarely tested. We examine a potential function of PC third‐party affiliation (affiliation between former opponents and bystanders) in rooks and jackdaws by investigating the hypothesis that conflicts lead to further aggression and that PC third‐party affiliation increases to reduce such aggression. The results show that PC affiliation reduces PC aggression for rook aggressors who were less likely to receive aggression after conflicts when they were affiliating with another vs. when they were alone. The opposite result was found for victims of both species who received more aggression after conflicts, and this aggression was not reduced by the act of affiliating. Finally, for jackdaw aggressors, the amount of aggression received after conflicts was not influenced by whether the individual was affiliating or alone, indicating that PC third‐party affiliation may serve a function that we did not examine. These findings highlight the importance of investigating functional differences in PC affiliative behavior according to the role played in the conflict.     

Experimental Manipulation of Food Accessibility Affects Conflict Management Behaviour in Ravens

Conflict management strategies such as reconciliation and bystander affiliation have been described for a variety of species. A common determinant seems to be a ‘complex’ social life, with individuals relying on affiliate relationships or social bonds. Little is known, however, about the strategic and flexible use of conflict management skills in experimental settings in species other than primates. We here investigated conflict and post‐conflict behaviour of ravens by manipulating the accessibility of food and, thus, the likelihood of aggressive interactions while foraging. Specifically, we presented birds with a certain amount of highly preferred food that varied in the number of pieces (one piece, two pieces or, as a control, small pieces matching the number of participating birds) and observed their agonistic behaviour during feeding and their affiliative behaviour afterwards. The results showed that high levels of conflicts during feeding in the 1‐piece and 2‐piece conditions led to high levels of affiliation after feeding. Depending on the experimental condition, this effect is best explained (a) by the affiliative behaviour of former aggressors (1‐piece condition) and (b) by the affiliation directed to the receivers of aggression after feeding (2‐piece condition). Those dyads that engaged in allo‐preening after feeding also engaged in allo‐preening outside the experimental setting, suggesting that socially bonded individuals provided third‐party affiliation to victims of aggression. Moreover, socially bonded ravens fed close to each other in the experiment when food was clumped, indicating that they actively coordinated their behaviour when there was a high conflict potential. Taken together, these findings support the assumption that ravens use their social bonds to avoid conflicts by choosing with whom to feed, and to buffer effects of conflicts by engaging in third‐party affiliation as post‐conflict behaviour.     

Post-conflict Behaviour of Wild Olive Baboons. I. Reconciliation,Redirection and Consolation

Observations of post-conflict interactions have provided important insights into primate social organization. In this study, the nature and determinants of post-conflict behaviour in a troop of wild olive baboons, Papio anubis, were investigated. Reconciliation was observed among all age-sex classes, occurring at a rate consistent with a relatively intolerant dominance style. Reconciliation was more frequent when one of the combatants carried a dependent infant but rarely followed conflicts associated with food. Neither the directionality nor the decidedness of conflicts affected conciliatory tendency. In contrast, opponents that were close kin or of similar rank reconciled more often. Olive baboons did not affiliate with non-combatants more frequently following aggression than in control periods, although affiliation with supporters and the close kin of opponents increased. Absence of consolation follows the observed cercopithecine pattern, consistent with the hypothesis that consolation requires an ability to empathize with the victim's distress. Initiation of post-conflict attacks on third parties was not elevated in victims of aggression. The rarity of redirection is attributed to spatial dispersion, the frequent bidirectionality of baboon aggression and regular male intervention in female conflicts, all of which appear to limit the availability of ‘safe’ targets.     

Reconciliation following aggression in patas monkeys,Erythrocebus patas

Following intra-group aggression, obvious conciliatory displays are absent from the behavioural repertoire of patas monkeys, Erythrocebus patas, while many other Old World primate species show special reconciliation gestures. When 10-min focal-animal samples that began after spontaneous aggression were compared with matched-control samples, captive adult female patas followed up on aggressive interactions, interacting sooner and more often with former opponents during post-conflict observations than during matched-control observations. Almost one-third of post-conflict observations included affiliative behaviour between former opponents, which is termed reconciliation. Matrilineally related opponents were more likely to reconcile with one another than were unrelated animals. No effect of the dominance hierarchy on tendency to reconcile was found. Thus, patas monkeys showed general patterns in post-conflict behaviour that were similar to those seen in other primates previously investigated.     

Reconciliation in the Spotted Hyena (Crocuta crocuta)

Theory predicts that it should often be in the best interests of gregarious animals to repair social bonds damaged by within-group conflict. Indeed, reconciliation in many primates takes the form of affiliative behavior occurring shortly after a conflict. Here we inquired whether reconciliation also occurs among spotted hyenas ( Crocuta crocuta ), gregarious carnivores whose social lives share much in common with those of cercopithecine primates. In a large group of free-living hyenas in Kenya, we used focal animal observations to monitor rates at which various affiliative behaviors occurred before and after dyadic aggressive interactions. An affiliative behavior was identified as having a conciliatory function only if it occurred more frequently after than before fights, and if it was also associated with reduced rates of aggression between former opponents during the post-conflict interval. Of all affiliative behaviors monitored, only two types satisfied both these criteria: greeting behavior and non-aggressive approach. Over 72% of conciliatory behaviors occurred within the first five minutes after a fight, and hyenas reconciled after 14.6% of 698 fights. Mean conciliatory tendency (CT) for individual hyenas was 11.3%. Hyenas exhibited higher CTs when they were recipients (victims) of aggression than when they were aggressors, and they showed higher CTs in interactions with non-kin than with kin. Conciliatory tendencies did not vary with age–sex classes of opponents or with rank distance between opponents . Conciliatory tendency in spotted hyenas fell at the low end of the CT range observed among primates.     

Duration and outcome of intergroup conflict influences intragroup affiliative behaviour

Theoreticians have long suggested that the amount of intergroup conflict in which a group is involved could influence the level of cooperation or affiliation displayed by its members. Despite the prevalence of intergroup conflicts in many social animal species, however, few empirical studies have investigated this potential link. Here, I show that intragroup allopreening rates are highest in green woodhoopoe (Phoeniculus purpureus) groups that have the greatest involvement in intergroup conflict. One reason for this relationship is a post-conflict increase in allopreening, and I demonstrate for the first time that both conflict duration and outcome influence subsequent allopreening rates: group members allopreened more following long conflicts and those they lost compared with short conflicts and those they won, perhaps because the former are more stressful. The increase in affiliative behaviour was the result of more allopreening of subordinate helpers by the dominant breeding pair, which may be because the breeders are trying to encourage helpers to participate in future conflicts; relative group size influences conflict outcome and helpers contribute more to conflicts than do the breeding pair. These results emphasize that our understanding of cooperation and group dynamics can be enhanced by investigations of how intergroup interactions affect intragroup processes.     

Anxiety Level Predicts Post-Conflict Behaviour in Wild Japanese Macaques (Macaca fuscata yakui)

Reconciliation (i.e. the post-conflict exchange of friendly behaviour between former opponents) functions to control for the detrimental effects that aggression may have on social relationships. Studies conducted so far have investigated intra-individual sources of variation in post-conflict behaviour, showing that animals have a stronger increase in anxiety and are more likely to reconcile after conflicts with valuable partners, such as kin. Much less attention has been given to how differences between individuals in emotional profiles affect post-conflict behaviour. Our aim was to analyse whether inter-individual differences in baseline anxiety levels predicted the magnitude of the increase in anxiety following a conflict and the occurrence of reconciliation. We collected data on two groups of wild Japanese macaques ( Macaca fuscata yakui ). Animals having a higher baseline level of anxiety had a more dramatic anxious response following a conflict while controlling for a series of factors (e.g. relationship quality between opponents). These more anxious animals were also less likely to reconcile than more relaxed individuals. Therefore, more anxious animals face some social costs by being less able to cope with the post-conflict condition. We propose that differences in anxiety levels may be interpreted as tradeoffs between benefits and costs across conditions. For example, more anxious animals, who are less able to reconcile conflicts, might also be less exploratory and thus face a lower risk to eat unknown, poisonous food.     

Canine Reconciliation and Third‐Party‐Initiated Postconflict Affiliation: Do Peacemaking Social Mechanisms in Dogs Rival Those of Higher Primates?

Conflicts over food, access to mates, or other limited resources can sometimes escalate into aggression. In species that form social groups, these aggressive conflicts can jeopardize the benefits of group living, such as enhanced access to valued resources, necessitating the development of behavioural mechanisms that either mitigate conflicts, prevent aggressive escalation or resolve disputes. Two important mechanisms for managing the effects of disputes involve postconflict (PC) affiliative behaviour, either between the former opponents (called ‘reconciliation’) or between one opponent and a third party. Even though numerous studies have tested reconciliatory and third‐party affiliation tendencies in primates, for non‐primate animals little systematic data are available. We performed behavioural observations on three groups of captive domestic dogs (Canis familiaris) and used a PC/MC method to test the existence of reconciliation and third‐party affiliation in this species. The results of this study clearly indicate that both reconciliation and third‐party PC interactions are present in the domestic dog, and form important social mechanisms of the domestic dog. Furthermore, familiar individuals showed a significantly higher proportion of reconciled conflicts than did unfamiliar individuals, and also displayed fewer conflicts. Finally, we show that most of the third‐party affiliations involve the victim of a conflict and that victim‐directed affiliation outweighs victim‐initiated third‐party affiliation in these PC interactions.     

Patterns of reconciliation among captive gelada baboons (Theropithecus gelada): A brief report

Animals that live in groups are frequently exposed to conflict situations and must in some way maintain group cohesion. One mechanism that appears to restore social relationships after they have been disrupted by conflict isreconciliation. This study investigated reconciliatory behavior in the gelada baboon,Theropithecus gelada. The subjects were 11 adult geladas, housed in a large outdoor enclosure at the Bronx Zoo/Wildlife Conservation Park, New York. Five-minute focal animal samples following spontaneous aggression were compared with 5-min matched-control samples. The results of this study were: (1) geladas reunited in a friendly way after aggression; (2) former opponents were attracted to one another rather than dispersed from one another after a conflict; (3) most post-conflict reunions occurred within the first 2 min of the post-conflict period; and (4) geladas do not have any specific types of behavior associated with post-conflict reunions as do chimpanzees and macaques. The results of this study support the hypothesis that gelada baboons reconcile after aggression.     

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.     

Postconflict third-party affiliation in rooks, Corvus frugilegus

Conflict features in the lives of many animal species and induces social stress mediated by glucocorticoid hormones [1]. Postconflict affiliation, between former opponents (reconciliation) or between former opponents and a bystander (third-party affiliation), has been suggested as a behavioral mechanism for reducing such stress [2], but has been studied almost exclusively in primates [3]. As with many primates, several bird species live in social groups and form affiliative relationships [4]. Do these distantly related animals also use affiliative behavior to offset the costs of conflict? We studied postconflict affiliation in a captive group of rooks. Unlike polygamous primates, monogamous rooks did not reconcile with former opponents. However, we found clear evidence of third-party affiliation after conflicts. Both initiators and targets of aggression engaged in third-party affiliation with a social partner and employed a specific behavior, bill twining, during the postconflict period. Both former aggressors and uninvolved third parties initiated affiliative contacts. Despite the long history of evolutionary divergence, the pattern of third-party affiliation in rooks is strikingly similar to that observed in tolerant primate species. Furthermore, the absence of reconciliation in rooks makes sense in light of the species differences in social systems.     

A Statistical Modelling Approach to the Occurrence and Timing of Reconciliation in Wild Japanese Macaques

In various social species, animals have been observed to share friendly relationships with some group members and to resolve conflicts through reconciliation, the exchange of affiliative behaviour soon after a conflict that functions to restore the relationship between the former opponents. The valuable relationship hypothesis predicts that reconciliation should be observed more often after conflicts between friends. Friendly relationships can be described by three dimensions (i.e. value, security and compatibility); however, research into the relative importance of these dimensions for the occurrence of reconciliation is sparse. Moreover, reconciliation may depend on factors other than the social relationship between opponents including, for example, their social status or the context of the conflict. Our study aimed at analysing which factors are important determinants of reconciliation and at testing the valuable relationship hypothesis, by analysing the relative effects of relationship value, security and compatibility on the occurrence and timing of reconciliation. We collected data on two troops of wild Japanese macaques living on Yakushima Island, Japan, and selected the best predicting variables of reconciliation using linear mixed models. Our results show that reconciliation occurs more frequently, and earlier, after conflicts between opponents who exchange a higher percentage of grooming. Two additional variables related to relationship security and value were selected in the best models: frequency of aggression and of approaches resulting in tolerated co‐feeding. Among the variables not related to relationship quality, distance between opponents at the end of the conflict, kinship, sex of the opponents and context of conflict (i.e. during feeding or social time) were included in our models. Our findings support the valuable relationship hypothesis and, in particular, highlight that the fitness‐related benefits of social relationships (i.e. the relationship value) are important determinants of the evolution of friendly relationships and reconciliation.     

Effect of valuable relationship on reconciliation and initiator of reconciliation in captive bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>)

Reconciliation, post-conflict affiliation between former opponents, was suggested to repair the relationship between former opponents. The number of times that each former opponent initiates reconciliation may be affected by the dominance style. Despotic dominance style was suggested to restrain the initiation of reconciliation by recipients of aggression, unlike relaxed dominance style. Although reconciliation was suggested to restore relationships, reconciliation sometimes did not occur. The valuable relationship hypothesis predicts that reconciliation is more likely to occur when former opponents share a valuable relationship. However, few studies have tested this hypothesis in non-primate species. This study investigated, in captive bottlenose dolphins, which are suggested as having a relaxed dominance style, whether aggressors or recipients of aggression initiate reconciliation, and whether relationships between former opponents affect the occurrence of reconciliation. The number of times that aggressors or recipients of aggression initiate reconciliation did not differ. Recipients of aggression initiated reconciliation sooner than aggressors. These results support the prospect that recipients of aggression are not impeded in initiating reconciliation in relaxed species. Former opponents who share a more affiliative relationship were reconciled more frequently. Bottlenose dolphins were suggested to receive the benefits from individuals who shared more affiliative relationships. The valuable relationship hypothesis may be supported in bottlenose dolphins.